Henderson, L. (2007). Invasive, Naturalized and Casual Alien Plants in Southern Africa

Home , Acacia mearnsii, Limonium, Myoporum tenuifolium, Tagetes minuta, Triplaris americana

Bothalia 37,2: 215–248 (2007) Invasive, naturalized and casual alien plants in southern Africa: a summary based on the Southern African Plant Invaders Atlas (SAPIA)

L. HENDERSON*

Keywords: biomes, casual alien plants, invasive plants, Lesotho, naturalized plants, roadside surveys, SAPIA mapping project, South Africa, Swaziland

ABSTRACT

The primary objective of this publication is to provide an overview of the species identity, invasion status, geographical extent, and abundance of alien plants in South Africa, Swaziland and Lesotho, based on ﬁ eld records from 1979 to the end of 2000. The dataset is all the species records for the study area in the Southern African Plant Invaders Atlas (SAPIA) database during this time period. A total of 548 naturalized and casual alien plant species were catalogued and invasion was recorded almost throughout the study area. Most invasion, in terms of both species numbers and total species abundance, was recorded along the southern, southwestern and eastern coastal belts and in the adjacent interior. This area includes the whole of the Fynbos and Forest Biomes, and the moister eastern parts of the Grassland and Savanna Biomes. This study reinforces previous studies that the Fynbos Biome is the most extensively invaded vegetation type in South Africa but it also shows that parts of Savanna and Grassland are as heavily invaded as parts of the Fynbos. The Fabaceae is prominent in all biomes and Acacia with 17 listed species, accounts for a very large proportion of all invasion. Acacia mearnsii was by far the most prominent invasive species in the study area, followed by A. saligna, Lantana camara, A. cyclops, Opuntia ﬁ cus-indica, Solanum mauritianum, Populus alba/×canescens, Melia azedarach, A. dealbata and species of Prosopis.

INTRODUCTION transects of between fi ve and 10 km long. Recordings of streambank species were made at virtually all water- History of roadside surveys in South Africa course crossings on the survey route. Roadside surveys of invasive plants in South Africa The Southern African Plant Invaders Atlas mapping were pioneered by Henderson and Musil (née Duggan) project (SAPIA) starting in 1979 in the central Transvaal, now Gauteng (Wells, Duggan & Henderson 1980), with the remainder The Southern African Plant Invaders Atlas (SAPIA) is of the Transvaal surveyed in 1982 and 1983 (Henderson a mapping project, launched in January 1994, to collate & Musil 1984). Surveys of the rest of South Africa were information on the distribution, abundance and habitat conducted by Henderson from 1986, starting with Natal types of invasive and naturalized alien plants in southern (Henderson 1989), followed by the Orange Free State Africa (Henderson 1998b). The fi rst phase of SAPIA, (Henderson 1991a), northern Cape (Henderson 1991b), involving volunteer participants, was scheduled for a eastern Cape (Henderson 1992), western and central fi ve-year period, ending in December 1998. The atlas Cape (completed in 1993 but unpublished), and southern region covered South Africa, Lesotho and Swaziland. and southwestern Cape (Henderson 1998a). Information was recorded on two standardized atlas All terminology used in this paper relating to invasive sheets, with slightly different species lists, covering the plants such as ‘alien’, ‘invasive’, ‘naturalized’, ‘casual western and eastern halves of the atlas region. One hun- alien’, ‘weed’ and ‘environmental weed’ conforms, as dred plant taxa were listed on each sheet, with a com- far as possible, to the defi nitions provided by Richardson bined total of 161 species. A pocket fi eld guide was com- et al. (2000) and Pyšek et al. (2004). The method used piled to help with the identifi cation of all listed species in these surveys was designed initially to make use of (Henderson 1995). otherwise unproductive travelling time whilst engaged in other research projects. The method was refi ned as SAPIA database the surveys progressed until a standardized method was developed (see Henderson 1992, 1998a). The presence A computerized SAPIA database was created by and abundance of all alien trees, large shrubs and con- incorporating all Henderson survey data (± 23 000 spicuous climbers which appeared to be naturalized or records) and SAPIA phase one project data (± 20 000 occurring outside of cultivation were recorded for each records). The SAPIA project continued on an ad hoc veld type category, habitat type (roadsides and adjoining basis and by the end of 2000 a total of ± 48 000 records veld, and streambanks) and quarter-degree/fi fteen minute had been accumulated. Thereafter, the SAPIA initiative square traversed by road. dwindled due to lack of funding. Only 10 000 records were added in the fi ve year period from 2001 to the Recordings of species on roadsides and in the adja- end of 2005. The SAPIA project was revived in 2006 cent veld were made from a moving vehicle along road with funding from the Department of Water Affairs and Forestry’s Working for Water Programme. The SAPIA * Agricultural Research Council: Plant Protection Research Institute, c/o SANBI, Private Bag X101, 0001 Pretoria. database has been computerized using Microsoft Access E-mail: [email protected] and is housed at the Plant Protection Research Institute MS. received: 2006-10-06. in Pretoria. 216 Bothalia 37,2 (2007) Objectives of this study Data treatment • To provide an overview of the species identity, inva- Abundance sion status, geographical extent, and abundance of alien plants in South Africa, Swaziland and Lesotho, Species abundance ratings in the SAPIA database are based on fi eld records from 1979 to the end of 2000. qualitative estimates. Table 1 shows the abundance ratings used in the SAPIA database and the equivalent rat- • To highlight the most prominent invaders in the ing used in Henderson surveys. For the purposes of this region as a whole, in each of the biomes, and in study, species abundance ratings were converted to a riparian and wetland habitats. numerical value as done in previous surveys (Henderson • To compare invasion and provide species profi les for 1998a) and each abundance rating was expressed in each of the biomes. numbers of individuals or groups per 10 km transect/ recording (Table 1).

METHODS Prominence A similar formula was used in this study to calcu- Sampling method late prominence as in previous studies by Henderson The dataset for this study is all the species records for (1998a). The prominence value of a species x in category South Africa, Swaziland and Lesotho in the SAPIA data- y (biome or study area) was calculated as follows: base collected from 1979 until the end of 2000 (± 48 000 total abundance of species x in category y records). During this period a concerted effort was made × 100 to gather as much data from as wide an area as possi- sum of the abundances of all species in category y ble. The information gathered is the best available data prominence = + concerning the extent of invasion and species composi- value tion, at least of the larger trees, shrubs and conspicuous total species records of species x in category y climbers, in the study area over this time period. × 100 sum of the records of all species in category y The SAPIA dataset was subdivided on a quarter- degree square (QDS) basis into six datasets representing The highest prominence values in a given category the biomes of southern Africa. According to Rutherford which add up to ± 160 points out of a total of 200 are (1997) there are seven biomes in southern Africa: Savanna, printed in bold in Appendices 1–3. The cut-off point is Fynbos, Forest, Grassland, Nama-Karoo, Succulent Karoo arbitrary but represents the upper 80% of the summed and Desert. The Forest Biome in southern Africa is min- prominence values. iscule, only occurring in the Knysna area. However, if all the forest patches elsewhere are included, its area increases several-fold (Rutherford 1997). In this study RESULTS Forest refers to the Forest Biome and also forest habitats within the Savanna, Fynbos and Grassland Biomes. The A total of 548 naturalized and casual alien plant spe- Desert Biome occurs almost exclusively in Namibia, cies were catalogued in the SAPIA database for South except for a very small patch along the Orange River Africa, Swaziland and Lesotho from 1979 to the end of bordering on South Africa that has been excluded from 2000 (Appendix 4). At least 119, mainly herbaceous, this study. taxa are considered to have been under-recorded and

TABLE 1.—Abundance ratings used in Henderson surveys, SAPIA and this study

Henderson surveys SAPIA This study Rating Roadsides and veld No.* Streambanks Rating All habitats† All habitats# 9 A virtually continuous, almost pure stand 1 000+ Any number, with cover more than 7 Very abundant 1 000 75% of the reference area 8 The commonest species in a generally con- 500–999 Any number, with 50–75% cover 6 Very abundant 1 000 tinuous tree or shrub layer 7 Less abundant than above but > 20 individu- 200–499 Any number, with 25–50% cover 5 Abundant 200 als or groups per km 6 10–20 individuals or groups per km 100–199 Any number with 5–25% cover 4 Abundant 200 5 5–10 individuals or groups per km 50–99 Numerous, but < 5% cover or scat- 3 Frequent 50 tered, with cover up to 5% 4 2–5 individuals or groups per km 20–49 Few, with small cover 2 Frequent 50 3 ± 1 individual or group per km 5–19 Solitary, with small cover 1 Occasional 10 2 Less abundant than above but more than 1 2–4 Occasional 10 individual or group per 5 km 1 ± 1 plant or group per 5–10 km 1 Rare 1 *, approximate numbers of individuals or groups per 10 km transect. †, very abundant extensive stands; abundant; many clumps or stands: frequent, many sightings of single plants or small groups: occasional, a few sightings of one or a few plants: rare, one sighting of one or a few plants. #, weighted abundance, numbers of individuals or groups per 10 km transect/recording. Bothalia 37,2 (2007) 217 the results presented are not a true reﬂ ection of their status (see asterisked species in Appendix 4). A further 45 species were recorded in the study area after 2000 and are asterisked in the species checklist (Appendix 5). A total of 601 species are listed in the full checklist given in Appendix 5—this is estimated to be about half the total number of naturalized and casual alien plant species in southern Africa. The most comprehensive listing of naturalized species in southern Africa, compiled by Wells et al. (1986), contains approximately 965 species, predominantly herbaceous. The SAPIA database, with a bias towards trees and shrubs, has an additional 231 species not listed by Wells et al. (1986).

Geographical extent of invasion A Alien plant invasion was recorded almost throughout the study area. Figure 1A shows invasion in terms of species numbers per QDS and Figure 1B shows the severity of invasion per QDS based on the total weighted abundance of all species per QDS. Most invasion, in terms of both species numbers and total species abundance, was recorded along the southern, southwestern and eastern coastal belts and in the adjacent interior. This corresponds with the regions of highest rainfall (Schulze 1997), urban development, and cultivation of agricultural and silvicultural crops. It also includes the whole of the Fynbos and Forest, and the moister eastern parts of the Grassland and Savanna Biomes (Figure 1C). Distribution maps of 234 species, which include all declared species under the Conservation of Agricultural Resources Act, Act 43 of 1983, and amended in 2001, B are given in the ﬁ eld guide Alien weeds and invasive plants (Henderson 2001).

Prominent invasive species There were 97 prominent invasive species in the study area and each of the biomes (Appendices 1–3). All these species were invading natural and semi-natural habitats. Study area Fifty species account for most invasion (the upper 80% of the summed prominence values) in the study area (Appendix 1). Acacia mearnsii (black wattle) was the most prominent species by far, with a value of 18.37 (out of a maximum of 200) which is more than double the value of the second-ranked species, A. saligna (Port C Jackson). The remaining top ten most prominent invaders in the study area were in order, Lantana camara (lantana), A. cyclops (rooikrans), Opuntia fi cus-indica FIGURE 1.—A, species numbers per quarter-degree square in study (sweet prickly-pear), Solanum mauritianum (bugweed), area; B, severity of invasion per quarter-degree square. Light Populus alba/×canescens (white/grey poplars—values invasion: < 1 individual or group per km. Moderate invasion: up to 5 individuals or groups per km; some species forming stands. of these two taxa were combined where they were diffi - Heavy invasion: up to 50 individuals or groups per km; many cult to distinguish at a distance during roadside surveys), species forming stands; some completely dominating landscape. Melia azedarach (seringa), A. dealbata (silver wattle) C, heavy invasion in relation to biomes in study area. and Prosopis spp. (P. glandulosa var. torreyana, P. velutina and their hybrids)(mesquite trees). Together these was the most prominent species with a prominence value species cover almost the entire study area (Figures 2, 3). of 20.6, followed by Chromolaena odorata (triffi d weed) Savanna Biome with a value of 14.2 and Melia azedarach with a value of 12. The remaining top ten invaders were, in order, Forty-eight species were the most prominent invaders Solanum mauritianum, Acacia mearnsii, Opuntia fi cus- in the Savanna Biome (Appendix 2). Lantana camara indica, Ricinus communis (castor-oil plant), Psidium 218 Bothalia 37,2 (2007)

A B

C D

E F

FIGURE 2.—Distribution and severity of invasion in study area: A, Acacia cyclops; B, Acacia dealbata; C, Acacia mearnsii; D, Acacia saligna; E, Lantana camara; F, Melia azedarach. Light invasion, U; moderate invasion, S; heavy invasion, . guajava (guava), Eichhornia crassipes (water hyacinth) blackwood), A. longifolia (long-leaved wattle), Populus and Jacaranda mimosifolia (jacaranda). ×canescens (grey poplar), Paraserianthes lophantha (stinkbean), Rubus fruticosus (European blackberry) and Fynbos Biome Opuntia ﬁ cus-indica. Hakea sericea (silky hakea) and Twenty species were the most prominent invaders in Pinus radiata (radiata pine), both invaders of mountain the Fynbos Biome (Appendix 2). Acacia mearnsii was fynbos, were most likely under-recorded because of the the most prominent species with a prominence value of inacessibility and under-sampling of this habitat. 31.5, followed by A. saligna and A. cyclops with val- Forest habitats ues of 30.4 and 27.2, respectively. The remaining top ten most prominent invaders in order, were, Pinus pin- Forty species were the most prominent invaders in aster (cluster pine), Acacia melanoxylon (Australian forest habitats (Appendix 2). Chromolaena odorata was Bothalia 37,2 (2007) 219

A B

C D

FIGURE 3.—Distribution and severity of invasion in study area: A, Opuntia fi cus-indica; B, Populus alba/×canescens; C, Prosopis spp.; D, Solanum mauritianum. Light invasion, U; moderate invasion, S; heavy invasion, . the most prominent species with a prominence value 2000 and currently would be rated as one of the most of 23.9, followed by Solanum mauritianum and Acacia prominent invaders in the Grassland Biome (Henderson mearnsii with values of 19 and 16.7, respectively. The et al. 2003). remaining top ten prominent invaders were, in order, Acacia melanoxylon, Lantana camara, Cestrum laevi- Nama-Karoo Biome gatum (inkberry), Caesalpinia decapetala (Mauritius/ Mysore thorn), Melia azedarach, Pinus pinaster and Fourteen species were the most prominent invaders in Psidium guajava. Pereskia aculeata (pereskia) ranked the Nama-Karoo Biome (Appendix 3). Prosopis spp. (P. eleventh and could have been vastly underestimated glandulosa var. torreyana, P. velutina and their hybrids) because of the diffi culty of observing this forest canopy, were the most prominent species with a prominence climbing species. value of 60.6, followed by Atriplex infl ata (sponge-fruit saltbush) and Opuntia fi cus-indica with values of 21 and Grassland Biome 14 respectively. The remaining top ten prominent invaders were, in order, Salsola kali/tragus (Russian tumble- Thirty-two species were the most prominent invaders weed), Azolla fi liculoides (red water fern), Nicotiana in the Grassland Biome (Appendix 3). Acacia mearnsii glauca (wild tobacco), Atriplex nummularia (old man was the most prominent species with a prominence value saltbush), Schinus molle (pepper tree), Agave americana of 21.3, followed by A. dealbata and Salix babylonica (American agave) and Solanum elaeagnifolium (silver- (weeping willow) with values of 20.9 and 17.3, respec- leaf bitter-apple). tively. The remaining top ten most prominent invaders were, in order, Populus alba/×canescens (white/grey Succulent Karoo Biome poplars), Solanum mauritianum, Rubus spp. (mainly R. cuneifolius)(brambles), Pyracantha angustifolia and P. Twelve species were the most prominent invaders in crenulata (yellow and Himalayan fi re thorns), Eucalyptus the Succulent Karoo Biome (Appendix 3). Nicotiana spp. (eucalypts), Melia azedarach and Opuntia fi cus- glauca was the most prominent invader with a promi- indica. Campuloclinium macrocephalum (pompom nence value of 26.8, followed by Acacia cyclops and weed) which did not feature as a prominent invader in Prosopis spp. (P. glandulosa var. torreyana, P. velutina this study showed an explosive rate of increase after and their hybrids) with values of 26.3 and 25.9, respec- 220 Bothalia 37,2 (2007) tively. The remaining top ten most prominent invaders nent invasive species in each of the biomes and the study were, in order, Acacia mearnsii, A. saligna, Atriplex area in terms of region of origin, taxonomy, growth infl ata, Arundo donax (giant reed), Atriplex nummularia, form, perennation, type of reproduction, dispersal mecha- Opuntia fi cus-indica and Populus ×canescens. nism and cultivated use.

Riparian and wetland habitats Savanna Biome species are predominantly of tropical origin; members of the Fabaceae, Solanaceae, Asteraceae Fifty-ﬁ ve species had more than 50 records in ripar- and Rosaceae; woody trees and shrubs, followed by ian and wetland habitats (Appendix 4). Salix babylonica herbs and climbers; perennial evergreen and evergreen/ was the most frequently recorded riparian and wetland deciduous; seed-producers; water and bird dispersed; species with 1 323 records, followed by Populus alba/ ornamentals and agricultural crops. ×canescens with 1 176 records and Acacia mearnsii with 953 records. The remaining top ten riparian and wetland Fynbos Biome species are predominantly of temperate invaders were, in order, Melia azedarach, Ricinus com- origin (particularly southern temperate); members of the munis, Arundo donax, Acacia dealbata, Sesbania puni- Fabaceae, Myrtaceae, Pinaceae and Salicaceae; woody cea (red sesbania), Prosopis spp. and Nicotiana glauca. trees and shrubs; perennial evergreen; seed-producers; water, bird and wind dispersed; silvicultural crops, orna- Biome comparison mentals and cover/binders.

The Savanna Biome, which occupies the largest Forest habitat species are predominantly of tropical number of QDS (645) in the study area, had the great- origin; members of the Fabaceae, Asteraceae, Myrtaceae, est number of species (358) and the most invasion in Solanaceae, Pinaceae and Zingiberaceae; woody trees terms of total abundance of all species (Table 2). The and shrubs, followed by herbs and climbers; perennial Fynbos Biome, however, which occupies the least QDS evergreen; seed-producers; bird and water dispersed; (139), was the most heavily invaded in terms of average ornamentals, barriers and silvicultural crops. abundance of all species per QDS, average abundance of individual species per QDS and % QDS heavily invaded. Grassland Biome species are predominantly of north- The Grassland Biome ranks third after Fynbos for total ern temperate origin and the tropics; members of the abundance of all species, followed by Forest, Nama- Rosaceae, Fabaceae and Salicaceae; woody trees and Karoo and the Succulent Karoo Biome was the least shrubs, followed by herbs; perennial evergreen/decidu- invaded. ous and deciduous; seed-producers, but a greater percentage of species coppice and sucker than in other veg- Biome proﬁ les etation categories; water and bird dispersed; barriers, ornamentals and agricultural crops. Appendix 6 provides species characteristics of the prominent invasive species. Table 3 analyses the promi- Nama-Karoo Biome species are predominantly of northern temperate origin and the tropics; members of TABLE 2.—Biome comparison in terms of extent, numbers and abun- the Chenopodiaceae, Salicaceae, Cactaceae, Fabaceae, dance of species and severity of invasion Solanaceae and Tamaricaceae; woody trees and shrubs, followed by herbs and succulent trees and shrubs; peren- FB Fh SB GB NKB SKB nial evergreen/deciduous and deciduous; seed-producers, Extent in QDS 139 157 645 521 548 141 but a greater percentage of species reproduce by vegeta- Total species 216 172 358 319 105 69 tive division than in other vegetation categories; water Prominent invasive 20 40 48 32 14 12 and wind dispersed; agricultural crops and ornamentals. species Total abundance* 986 222 1165 811 211 67 524 653 419 895 723 589 Succulent Karoo Biome species are predominantly of Ave abundance per 7 098 1 417 1 808 1 558 386 479 temperate origin; members of the Fabaceae, Chenopo- QDS diaceae and Tamaricaceae; woody trees and shrubs; Ave abundance per 4 568 1 293 3 257 2 545 2 015 979 perennial evergreen and evergreen/deciduous; seed-pro- species ducers and reproduce vegetatively by coppicing; water % QDS light# 17 15 51 49 51 66 and wind dispersed; agricultural crops, ornamentals and % QDS moderate# 30 44 20 33 13 9 cover/binders. % QDS heavy# 47 41891 1 QDS, quarter-degree squares in Fynbos, Savanna, Grassland, Nama- Karoo and Succulent Karoo according to Rutherford (1997); QDS in forest habitats according to SAPIA database. DISCUSSION *, total weighted abundance of all species (see text). Prominent invasive species: species with highest prominence values Biome comparison: extent of invasion adding up to ± upper 80% of summed values (see text). #, % QDS lightly invaded: less than 1 individual or group per km; #, % QDS moderately invaded: up to 5 individuals or groups per km; No previous studies have enabled a direct comparison some species forming stands; #, % QDS heavily invaded: up to 50 of the extent of invasion in the different biomes using the individuals or groups per km; many species forming stands; some same parameters. This study reinforces previous studies completely dominating landscape. FB, Fynbos Biome; Fh, Forest habitats; SB, Savanna Biome; GB, that the Fynbos Biome is the most extensively invaded Grassland Biome; NKB, Nama-Karoo Biome; SKB, Succulent vegetation type in South Africa (Richardson et al. 1997) Karoo Biome. but it also shows that parts of Savanna and Grassland are Bothalia 37,2 (2007) 221

TABLE 3.—Analysis of region of origin, taxonomy, growth forms, perennation, reproduction, dispersal mechanisms and cultivated uses of prominent invasive species in each of the biomes, forest habitats and study area

Characteristics Savanna Fynbos Biome Forest habitats Grassland Nama-Karoo Succulent Karoo Study area (97 Biome (49 (24 spp.) (53 spp.) Biome (35 Biome (18 spp.) Biome (16 spp.) spp.) spp.) spp.) Region of origin Northern temperate 10 (20%) 7 (29%) 14 (26%) 18 (51%) 9 (50%) 6 (38%) 32 (33%) spp. Southern temperate 4 (8%) 10 (42%) 6 (11%) 3 (9%) 3 (17%) 5 (31%) 15 (15%) spp. Tropical spp. 34 (69%) 7 (29%) 33 (62%) 13 (37%) 6 (33%) 5 (31%) 49 (51%) Hybrid 1 (2%) 1 (3%) 1 (1%) Taxonomy Families 18 12 24 14 10 10 32 Families with 50% or Fabaceae (9); Fabaceae (8); Fabaceae (7); Rosaceae (9); Chenopodiaceae Fabaceae (5); Fabaceae (15); more of total species Solanaceae (7); Myrtaceae (3); Asteraceae (5); Fabaceae (6) (3); Salicaceae Chenopodiaceae (2); Rosaceae (9); (no. spp. in brackets) Asteraceae (5); Pinaceae (2); Myrtaceae (4); Salicaceae (6) (3); Cactaceae Tamaricaceae (2) Solanaceae (9); Rosaceae (4) Salicaceae (2) Solanaceae (4); (2); Fabaceae (2); Asteraceae (6); Pinaceae (3); Solanaceae (2); Salicaceae (6); Zingiberaceae (3) Tamaricaceae (2) Cactaceae (5); Myrtaceae (5) Growth form (spp.) Woody tree & shrub 26 (53%) 21 (88%) 33 (62%) 26 (74%) 9 (50%) 12 (75%) 60 (62%) Succulent tree & 5 (10%) 1 (4%) 1 (2%) 1 (3%) 3 (17%) 1 (6%) 6 (6%) shrub Climber 7 (14%) 0 8 (15%) 1 (3%) 0 0 9 (9%) Herbaceous 10 (20%) 1 (4%) 10 (19%) 6 (17%) 5 (28%) 2 (13%) 20 (21%) Grass/reed 1 (2%) 1 (4%) 1 (2%) 1 (3%) 1 (6%) 1 (6%) 2 (2%) Perennation (spp.) Perennial evergreen 20 (41%) 18 (75%) 33 (62%) 13 (37%) 6 (33%) 8 (50%) 51 (53%) Perennial evergreen/ 11 (22%) 3 (13%) 9 (17%) 8 (23%) 5 (28%) 5 (31%) 19 (20%) deciduous Perennial deciduous 7 (14%) 2 (8%) 5 (9%) 9 (26%) 5 (28%) 2 (13%) 14 (14%) Variable 3 (6%) 1 (4%) 2 (4%) 2 (6%) 1 (6%) 1 (6%) 3 (3%) Germinative (annual/ 8 (16%) 0 4 (8%) 3 (9%) 1 (6%) 0 10 (10%) biennial) Reproduction (spp.) by: Seeds/spores 44 (90%) 21 (88%) 50 (94%) 29 (83%) 14 (78%) 14 (88%) 91 (94%) Coppicing 23 (47%) 11 (46%) 25 (47%) 22 (63%) 8 (44%) 11 (69%) 44 (45%) Suckering 9 (18%) 3 (13%) 6 (11%) 9 (26%) 4 (22%) 2 (13%) 14 (14%) Division 6 (12%) 4 (17%) 3 (6%) 6 (17%) 5 (28%) 2 (13%) 10 (10%) Rhizomes 1 (2%) 1 (4%) 4 (8%) 1 (3%) 2 (11%) 1 (6%) 6 (6%) Stolons/runners 1 (2%) 0 3 (6%) 1 (3%) 0 0 3 (3%) Bulbils 1 (2%) 0 0 0 0 0 1 (1%) Dispersal (spp.) by: Wind 13 (27%) 7 (29%) 19 (36%) 4 (11%) 7 (39%) 6 (38%) 31 (32%) Water 27 (55%) 16 (66%) 22 (42%) 19 (54%) 10 (56%) 11 (69%) 47 (48%) Birds 17 (35%) 8 (33%) 25 (47%) 17 (49%) 5 (28%) 4 (25%) 40 (41%) Mammals 9 (18%) 5 (21%) 7 (13%) 3 (9%) 4 (22%) 5 (31%) 16 (16%) Humans 14 (29%) 5 (21%) 8 (15%) 10 (29%) 3 (17%) 3 (19%) 21 (22%) Ants 8 (16%) 3 (13%) 4 (8%) 6 (17%) 0 2 (13%) 10 (10%) Cultivated uses (spp.) Ornamental 20 (41%) 6 (25%) 21 (40%) 7 (20%) 5 (28%) 4 (25%) 35 (36%) Cover/binder 4 (8%) 5 (21%) 6 (11%) 4 (11%) 2 (11%) 3 (19%) 9 (9%) Barrier 5 (10%) 2 (8%) 9 (17%) 8 (23%) 2 (11%) 0 17 (18%) Silvicultural crop 2 (4%) 7 (29%) 8 (15%) 4 (11%) 0 2 (13%) 10 (10%) Agricultural crop 11 (22%) 4 (17%) 7 (13%) 7 (20%) 7 (39%) 7 (44%) 16 (16%) Species with no uses 7 (14%) 0 2 (4%) 5 (14%) 2 (11%) 0 11 (11%) 222 Bothalia 37,2 (2007) as heavily invaded as parts of the Fynbos. These fi nd- Africa and is the most widespread riverine invader, ings have important implications for the management occurring almost continuously from Louis Trichardt of alien plant invasions in South Africa. Without inter- in the Limpopo Province down the eastern seaboard to vention we can expect invasion to increase in all parts Cape Town, a distance of ± 2 500 km. Acacia cyclops of South Africa and particularly in the Grassland and stretches along almost the entire Cape coastline from Savanna Biomes where large areas are yet to be invaded Port Nolloth in the northwest to beyond East London in and many species are only starting to invade. the east, a distance exceeding 2 000 km. Acacia saligna stretches along the Cape coastline from Saldanha Bay Biome comparison: prominent invaders in the west to the Kei River in the east, a distance of ± 1 500 km. Each biome has a different suite of prominent invaders. In part, this can be explained by their pre-adap- Sixty-eight per cent of prominent invaders are peren- tion to the prevailing environmental conditions, but nial trees or shrubs. There are only two grasses listed as also to their history of planting. Most of these species prominent invaders and only 14 species as nonperennial were deliberately introduced and cultivated on a grand (annual, biennial or variable). Grasses and herbaceous scale as silvicultural and agricultural crops e.g. Acacia species are under-represented in the SAPIA database mearnsii, A. melanoxylon, Pinus pinaster and species of largely as a consequence of biassed recording of the Prosopis, as barriers e.g. Acacia dealbata, Hakea seri- larger, more conspicuous species. In southern Africa cea and Pyracantha angustifolia, as cover/binders e.g. the Poaceae is one of the largest plant families with Acacia cyclops, A. saligna and Populus ×canescens, and 847 indigenous species and 115 (12%) naturalized spe- ornamentals e.g. Melia azedarach and Lantana camara. cies (Gibbs Russell et al. 1990). However, only 30 grass species are listed in this publication. There is defi nitely Some species which have become prominent invad- a lack of expertise in identifying grasses in South Africa ers were not cultivated widely or on a grand scale e.g. and this is one of the reasons for the under-represen- Solanum mauritianum, Chromolaena odorata and tation of alien grasses in weed surveys. There is simi- Nicotiana glauca. Although the latter species have on larly an under-representation of the alien herbaceous occasion been cultivated as ornamentals they have man- Asteraceae. The South African National Biodiversity aged to disperse very effi ciently without human assis- Institute’s online species checklist at http://posa.sanbi. tance—C. odorata by wind, S. mauritianum by birds org/searchspp.php lists 125 alien herbaceous species in and N. glauca by wind, soil and water. South Africa, yet only 44 alien herbaceous species have been listed in this publication. Some species, although widely planted, have become prominent invaders in only one biome, indicating that Comparison with other studies environmental factors have limited their distribution. Examples are members of the family Rosaceae, such Versfeld et al. (1998) provide the only other assess- as Pyracantha angustifolia, P. crenulata, Cotoneaster ment of the extent and importance of invasive plants on franchetii and C. pannosus that are virtually restricted a national level. This study combined expert knowledge to high-altitude grasslands where it appears that freez- of local landowners and managers with existing data- ing winter temperatures are needed to trigger seed ger- bases such as those of provincial conservation authori- mination (Henderson 1989). Jacaranda mimosifolia is ties and national departments. The SAPIA database another species that has been planted throughout South was used as a means of data verifi cation particularly for Africa yet is only invasive in the moister parts of the areas where expert knowledge was lacking. Overall the Savanna and Forest Biomes. In its native northeastern assessment by Versfeld et al. (1998) relating to impor- Argentina, J. mimosifolia occurs mainly on river banks tance rankings and the distribution of dense infesta- under warmer-temperate, subhumid conditions (Poynton tions concurs with this study. Eight of the top ten invad- 1973)—environmental conditions which are similar to ing species or groups of species, ranked by condensed those in its naturalized range in southern Africa. A pre- invaded area, also appear within the top ten ranking in vious study by Henderson (2006b) showed that the cur- this study—these are: Acacia cyclops, Prosopis spp., A. rent distributions of invasive plants in southern Africa mearnsii, A. saligna, Solanum mauritianum, Opuntia are a refl ection of the climatic zones of their origin. spp., Melia azedarach and Lantana camara. Versfeld et al. (1998) include Pinus spp. and Hakea spp. within There are considerable differences in the species pro- the top ten ranking, whereas this study includes Populus fi les of the biomes but shared features are the promi- alba/×canescens and Acacia dealbata. The lower rank- nence of the family Fabaceae, woody trees and shrubs, ing of Pinus spp. and Hakea spp. in this study can be reproduction by seed and water dispersal. Within the explained by the under-sampling of mountain habitats, Fabaceae the Acacia species are the most numerous which are largely inaccessible by road, in which these with 17 listed species and account for a very large pro- species are invasive. portion of all plant invasion in South Africa. They are important invaders of all the major vegetation types Abundance data presented in this study suggests that except for those in the arid interior, where other legu- Versfeld et al. (1998) may have underestimated the minous invaders take over, namely species of Prosopis. area of invasion of Salix babylonica and Populus alba/ The most widespread and abundant acacias are Acacia ×canescens. In the present study these species were not mearnsii, A. cyclops and A. saligna. Acacia mearnsii has only the most frequently recorded invaders in riparian invaded the widest range of vegetation types in South and wetland habitats but their total weighted abundance Bothalia 37,2 (2007) 223 was in both instances more than Melia azedarch and merged aquatic plant that has invaded much of the USA Eucalyptus spp. (Appendix 1) which were rated above since the 1960s, and Chondrilla juncea (skeleton weed), Salix spp. and Populus spp. by Versfeld et al. (1998). a terrestrial herb that has become a major agricultural Other riparian species which may also have been under- weed in the USA, Canada and Australia. estimated include Arundo donax (giant reed), Morus alba (common mulberry) and Ricinus communis.

CONCLUSION Looking to the future The main objective of this paper was to provide a The Working for Water Programme (WfW) and biologi- historical overview of the extent and species composi- cal control tion of alien plant invasion in southern Africa from 1979 until the end of 2000. This snapshot of invasion will pro- Alien plant invasion is a dynamic process and there vide a yardstick by which we can measure our progress will undoubtedly be changes in species composi- or failure in the management of invasive alien plants in tion and prominence of invaders in the future. Many southern Africa. of the large tree species—mainly Acacia, Eucalyptus, Pinus, Populus, Prosopis species and Melia azedarach This publication will also contribute to the global have been targeted by a national clearing programme, knowledge of invasive alien plants. One of the most Working for Water (WfW), which started in October useful predictors of invasiveness is whether a species 1995 (Marais et al. 2004). To date there has not been is invasive elsewhere in the world (Richardson et al. an assessment of the affects of the WfW programme on 2004a). The lists of prominent invaders and other natu- the status of invasive alien infestations. The programme ralized species provided here will serve as a warning has been proposed for 20 years but Marais et al. (2004) to neighbouring countries and to those as far afi eld as indicate that even with the existing generous levels of Australia, New Zealand and the USA of potentially inva- funding, it is unlikely that the problem will be contained sive species in their regions. within the next half century. The results presented here are but a summary of the Biological control of invasive plants using introduced more than 50 000 records of invasive alien plants in insects and pathogens is the only sustainable, effec- the SAPIA database. Much more can be gleaned from tive and inexpensive solution to the most intractable of the SAPIA data. SAPIA has provided the raw data for the invasive alien plant problems (Marais et al. 2004). analyses that have been used to prioritize invasive alien When they are successful, the damage infl icted by bio- species for management (Robertson et al. 2003; Nel et logical control agents causes a decline in population al. 2004), to map the potential spread of invasive plants densities, distribution and, or, rates of spread of inva- (Rouget et al. 2004), to look at broad-scale distribution sive plants, and reduces the costs of other management patterns of invasive species (Richardson et al. 2004b), practices (Zimmermann et al. 2004). There have been to correlate patterns of alien plant species richness some outstanding successes with biocontrol in South with the environment and indigenous species richness Africa, dating back to the early and mid-1900s with (Richardson et al. 2005), to correlate patterns of invasion Opuntia monacantha (drooping prickly pear) and O. with interactions between environment, species traits and fi cus-indica, and in more recent years with several of human uses (Thuiller et al. 2006) and to look at potential the Acacia spp. (Zimmermann et al. 2004). Population range and residence time (Wilson et al. 2007). SAPIA monitoring of A. saligna in the Western Cape has shown has also played a crucial role in providing information marked decreases in population densities caused by the on invasive plants for the revision of the Conservation gall-forming rust fungus, Uromycladium tepperianum of Agricultural Resources Act, Act 43 of 1983, and the (Morris 1997; Wood & Morris 2007). drafting of the National Environmental Management: Biodiversity Act, Act 10 of 2004. New invaders Alien plant invasion is a dynamic process and there- Since 2000 a further 45 species have been added to fore it is essential that the SAPIA database be kept up-to- the SAPIA database for the study area (Appendix 5). date with current information. From October 2006 a sec- Another eight species, two of which are indigenous to ond phase of the SAPIA mapping project was launched South Africa, are naturalized in neighbouring Zimbabwe and all the SAPIA data will be available online at the and Malawi (Appendix 5). All but three of the additional Weeds and Invasive Plants (WIP) website, www.agis. species have been listed as weeds in A global compen- agric.za/wip (Henderson 2006a). dium of weeds (Randall 2002) and 28 species are environmental weeds elsewhere in the world and therefore have the potential to become invasive in South Africa. ACKNOWLEDGEMENTS Fourteen of the new species are ‘noxious weeds’ or restricted in California, Florida, Hawaii, New Zealand I thank all the people who have contributed in many and Australia—places with similar climates and with different ways to the compilation of the SAPIA data- which South Africa has many invasive species in com- base and have made it one of the most comprehensive mon. We should be especially wary of these species databases on invasive alien plants in southern Africa. which include some of the most damaging and costly Les Powrie and Mike Rutherford of the South African invaders such as Hydrilla verticillata (hydrilla), a sub- National Biodiversity Institute are thanked for the QDS 224 Bothalia 37,2 (2007) coverage of the biomes of South Africa which was used heritage. Tafelberg, Cape Town. to subdivide the SAPIA dataset into the six biome data- PYŠEK, P., RICHARDSON, D.M., REJMÁNEK, M., GRADY, L., sets for this publication. Special tribute is made to Mike WILLIAMSON, M. & KIRSCHNER, J. 2004. Alien plants in checklists and fl oras: towards better communication between Wells of the Botanical Research Institute for his inspi- taxonomists and ecologists. Taxon 53: 131–143. ration and mentorship during the development of road- RANDALL, R.P. 2002. A global compendium of weeds (Draft). side survey techniques from 1979 to the mid-1980s, Hawaiian ecosystems at risk. Available at www.hear.org/gcw. and to Helmuth Zimmermann of the Plant Protection RICHARDSON, D.M., MACDONALD, I.A.W., HOFFMANN, J.H. & Research Institute for his motivation and support of the HENDERSON, L. 1997. Alien plant invasions. In R.M. Cowling, D.M. Richardson & S.M. Pierce, Vegetation of southern Africa. SAPIA mapping project. SAPIA is an initiative of the Cambridge University Press, Cambridge. Agricultural Research Council (ARC): Plant Protection RICHARDSON, D.M., PYŠEK, P., REJMÁNEK, M., BARBOUR, Research Institute which has provided the infrastruc- M.G., PANETTA, F.D. & WEST, C.J. 2000. Naturalization and ture, basal funding and support since the mid 1980s to invasion of alien plants: concepts and defi nitions. Diversity and the present. External funding of SAPIA has been grate- Distributions 6: 93–107. RICHARDSON, D.M., MORAN, V.C., LE MAITRE, D.C., ROUGET, fully received from the Departments of Agriculture, M. & FOXCROFT, L.C. 2004a. Recent developments in the Environmental Affairs and Tourism, and Water Affairs science and management of invasive alien plants: connecting and Forestry (Working for Water Programme). the dots of research knowledge, and linking disciplinary boxes. South African Journal of Science 100: 126–128. RICHARDSON, D.M., ROUGET, M., HENDERSON, L. & NEL, J.L. 2004b. Invasive alien plants in South Africa: macroecological REFERENCES patterns, with special emphasis on the Cape Floristic region. In M. Arianoutsou & V. Papanastasis, Proceedings of the 10th GIBBS RUSSELL, G.E., WATSON, L., KOEKEMOER, M., SMOOK, MEDECOS Conference, April 25–May 1, 2004, Rhodes, Greece. L., BARKER, N.P., ANDERSON, H.M. & DALLWITZ, M.J. Millpress, Rotterdam. 1990. Grasses of southern Africa. 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APPENDIX 1.—Prominent invaders in study area

Scientifi c name QSp QSa Tr A Pv R Scientifi c name QSp QSa Tr A Pv R Acacia Lantana camara 247 116 2 111 140 496 8.92 3 cyclops 166 91 1 097 203 636 8.66 4 Leptospermum laevigatum 38 15 102 15 916 0.71 dealbata 256 115 1 079 133 146 6.45 9 Litsea glutinosa 8 3 10 2 713 0.11 decurrens 101 31 232 23 456 1.23 Macfadyena unguis-cati 22 9 52 9 531 0.41 longifolia 94 34 363 53 080 2.43 Melia azedarach 551 65 2 119 65 735 6.64 8 mearnsii 428 251 2 620 410 950 18.37 1 Morus alba 129 8 304 5 983 0.85 melanoxylon 134 29 482 43 926 2.40 Nephrolepis exaltata 13 1 19 459 0.06 pycnantha 35 15 135 13 864 1.77 Nicotiana glauca 383 14 957 22 132 2.76 saligna 158 93 1 030 219 223 9.00 2 Opuntia Achyranthes aspera 77 3 85 1 262 0.22 fi cus-indica 861 57 2 445 72 477 7.55 5 Agave robusta 225 2 337 3 244 0.83 americana 431 8 761 9 136 1.94 stricta 106 14 193 5 412 0.59 sisalana 170 12 293 8 924 0.91 Paraserianthes lophantha 54 9 286 20 042 1.24 Ageratum Passifl ora edulis 32 0 55 261 0.13 conyzoides 37 8 50 5 141 0.27 Pennisetum clandestinum 48 12 53 8 884 0.39 conyzoides/houstonianum 31 8 49 2 994 0.20 Pereskia aculeata 44 8 102 5 788 0.40 houstonianum 26 2 31 1 728 0.12 Pinus patula 85 13 238 11 636 0.88 Argemone pinaster 85 44 401 48 229 2.36 mexicana 27 5 36 2 134 0.14 radiata 70 15 206 9 243 0.73 ochroleuca 154 20 206 12 115 0.82 sp. 14 2 15 1 338 0.07 Populus alba 15 2 22 915 0.08 7 Arundo donax 371 82 855 50 158 3.41 alba/canescens 185 47 460 33 871 2.04 7 Atriplex ×canescens 371 130 939 87 397 4.74 7 infl ata 164 77 213 58 878 2.28 nummularia 172 18 333 10 248 1.04 Prosopis glandulosa 40 10 50 4 988 0.26 10 Azolla fi liculoides 194 92 354 47 220 2.23 glandulosa/velutina 390 78 1 107 92 751 5.27 10 Caesalpinia decapetala 127 41 413 33 868 1.94 velutina 48 6 53 3 108 0.21 10 Cardiospermum grandifl orum 43 12 57 4 905 0.28 Prunus persica 319 1 728 7 401 1.81 grandifl orum/ 16 5 20 2 427 0.12 Psidium guajava 160 50 732 55 791 3.31 halicacabum Pyracantha angustifolia 142 3 285 3 735 0.74 Pyracantha angustifolia/cre- 40 3 51 1 183 0.15 Cereus jamacaru 124 11 193 13 042 0.82 nulata Cestrum laevigatum 70 16 167 11 039 0.70 Ricinus communis 456 56 1 701 48 855 5.21 Chromolaena odorata 93 64 558 137 654 5.46 Robinia pseudoacacia 110 14 178 8 828 0.66 Cinnamomum camphora 10 1 19 352 0.05 Rosa rubiginosa 119 12 276 11 494 0.95 Cirsium vulgare 188 20 345 14 022 1.18 Rubus Datura cuneifolius 75 35 236 49 313 2.03 ferox 175 14 201 8 685 0.71 fruticosus 89 32 244 22 810 1.23 innoxia 29 3 36 1 897 0.14 pascuus 3 2 3 450 0.02 sp. 84 1 110 1 710 0.29 sp. 86 30 179 29 694 1.30 stramonium 286 24 373 12 932 1.21 ×proteus 4 3 4 650 0.03 Eichhornia crassipes 87 72 431 79 893 3.40 Salix Eucalyptus babylonica 475 89 1 381 85 116 5.63 camaldulensis 121 22 182 13 259 0.80 fragilis 75 24 176 15 710 0.87 diversicolor 49 8 153 5 562 0.50 grandis 100 16 190 14 475 0.86 Salsola kali/tragus 155 31 187 14 080 0.84 sp. 505 30 1 103 23 523 3.12 Schinus molle 231 2 407 5 355 1.05 Senna didymobotrya 139 29 339 16 342 1.24 Hakea sericea 77 17 230 15 959 0.99 Sesbania punicea 323 68 830 52 078 3.41 Ipomoea Solanum indica 23 3 27 740 0.08 elaeagnifolium 51 11 60 14 136 0.57 indica/purpurea 74 7 120 2 284 0.33 mauritianum 265 99 1 364 135 219 7.14 6 purpurea 37 3 46 1 801 0.16 seaforthianum 30 3 77 2 656 0.25 sp. 3 0 3 52 0.01 Tithonia diversifolia 49 5 123 4 085 0.39 Jacaranda mimosifolia 195 16 613 17 430 1.87 Xanthium strumarium 149 21 212 12 633 0.85 Combined taxa e.g. Ageratum conyzoides/houstonianum indicate uncertainty of identifi cation. QSp, quarter-degree squares present; QSa, quarter-degree squares abundant; Tr, total records; A, total weighted abundance (see text); Pv, prominence value (bold numbers: highest prominence values which add up to ± upper 80% of summed values—see text); R, ranking of top ten taxa (taxa that are diffi cult to distinguish are grouped together).

APPENDIX 2.—Prominent invaders in Savanna Biome, Fynbos Biome and Forest habitats

Savanna Biome Fynbos Biome Forest Habitats Scientiﬁ c name QSp QSa R A Pv QSp QSa R A Pv QSp QSa R A Pv Acacia cyclops 31 20 181 28 557 3.40 102 63 810 174 964 27.20 5 0 9 214 0.73 dealbata 39 13 132 12 207 1.74 7 3 23 4 887 0.76 12 3 12 3 361 2.36 decurrens 19 7 38 1 779 0.35 4 1 4 1 061 0.76 longifolia 27 7 66 9 183 1.14 47 24 264 39 533 7.09 13 3 13 1 642 1.65 mearnsii 134 59 710 75 210 10.20 88 70 871 210 388 31.50 36 22 91 22 825 16.70 melanoxylon 33 2 73 1 274 0.49 50 23 298 38 206 7.35 25 12 85 18 186 14.20 Combined taxa e.g. Ageratum conyzoides/houstonianum indicate uncertainty of identiﬁ cation. QSp, quarter-degree squares present; QSa, quarter- degree squares abundant; R, records; A, total weighted abundance (see text); Pv, prominence value (bold numbers: highest prominence values which add up to ± upper 80% of summed values—see text). 226 Bothalia 37,2 (2007)

APPENDIX 2.—Prominent invaders in Savanna Biome, Fynbos Biome and Forest habitats (cont.)

Savanna Biome Fynbos Biome Forest Habitats Scientifi c name QSp QSa R A Pv QSp QSa R A Pv QSp QSa R A Pv Acacia (cont.) pycnantha 3 1 4 203 0.04 32 14 131 13 661 2.92 saligna 27 11 96 12 170 1.55 105 74 860 200 582 30.40 4 2 8 1 313 1.15 Achyranthes aspera 40 3 44 1 078 0.32 5 0 5 5 0.06 10 2 12 468 1.05 Agave americana 146 3 267 3 986 1.75 39 0 84 427 1.02 2 0 2 51 0.16 sisalana 132 12 251 8 801 2.08 10 0 10 46 0.12 3 0 4 22 0.29 Ageratum conyzoides 32 8 44 4 979 0.66 1 0 1 1 0.07 conyzoides/houstonianum 27 6 45 2 583 0.46 1 0 1 1 0.01 12 3 13 726 1.24 houstonianum 22 2 26 1 625 0.28 8 2 8 1 261 1.13 Argemone mexicana 23 4 32 1 082 0.26 1 1 1 1 000 0.11 1 0 1 50 0.09 ochroleuca 86 13 119 7 190 1.24 2 1 2 201 0.04 1 0 1 50 0.09 sp. 6 2 7 1 272 0.15 Arundo donax 132 34 372 20 126 3.69 71 25 172 16 721 3.70 3 1 3 1 020 0.67 Atriplex infl ata 13 9 16 7 701 0.75 19 10 25 7 148 1.02 nummularia 11 3 18 2 336 0.30 24 1 45 851 0.61 Azolla fi liculoides 43 16 75 4 979 0.83 16 10 37 7 491 1.19 2 2 2 400 0.32 Caesalpinia decapetala 80 22 239 20 966 3.06 17 7 25 6 053 4.48 Cardiospermum grandifl orum 40 11 54 4 703 0.69 5 3 7 1 511 1.17 grandifl orum/halicacabum 14 5 18 2 416 0.30 4 2 4 1 251 0.84 Cereus jamacaru 100 11 169 12 857 1.99 6 0 6 6 0.07 1 0 1 10 0.07 Cestrum laevigatum 48 12 138 7 551 1.38 6 0 9 58 0.11 22 9 30 6 457 5.01 Chromolaena odorata 79 57 529 133 524 14.20 33 25 77 41 029 23.90 Cinnamomum camphora 7 1 13 346 0.10 2 0 3 3 0.04 7 1 14 347 1.14 Cirsium vulgare 43 4 68 1 708 0.51 8 0 8 75 0.10 Datura ferox 69 3 87 2 676 0.69 3 0 3 3 0.04 2 0 2 11 0.15 innoxia 23 1 30 1 435 0.28 1 0 1 50 0.02 1 0 1 50 0.09 sp. 34 0 41 581 0.27 1 1 1 200 0.03 stramonium 104 9 131 4 082 1.04 15 1 19 1 357 0.36 6 1 6 1 170 0.95 Eichhornia crassipes 46 44 279 60 302 6.64 20 7 46 5 591 1.10 Eucalyptus camaldulensis 23 4 36 1 993 0.36 41 13 81 6 318 1.59 3 2 3 2 050 1.13 diversicolor 1 0 1 1 0.01 44 7 148 5 349 2.27 12 4 28 1 897 2.82 grandis 54 8 111 9 827 1.43 1 0 1 10 0.01 15 2 19 974 1.77 sp. 122 8 299 7 242 2.20 72 10 147 4 838 2.21 8 2 17 687 1.51 Hakea sericea 7 2 14 537 0.12 61 14 204 14 344 3.84 3 0 3 61 0.24 Ipomoea indica 18 3 22 735 0.18 3 0 3 3 0.04 3 0 3 3 0.21 indica/purpurea 49 3 86 1 101 0.55 10 1 18 409 0.25 11 2 16 1 308 1.71 purpurea 18 1 23 410 0.16 7 2 10 1 271 0.25 7 1 9 1 102 1.13 sp. 2 0 2 51 0.01 Jacaranda mimosifolia 139 16 497 16 767 4.06 17 2 21 1 589 2.19 Lantana camara 162 90 1 843 126 418 20.60 25 2 60 3 462 1.05 40 28 63 17 136 12.10 Leptospermum laevigatum 3 3 6 701 0.09 35 12 96 15 215 2.66 2 0 3 21 0.22 Litsea glutinosa 8 3 10 2 713 0.29 4 2 6 2 451 1.52 Macfadyena unguis-cati 17 6 47 8 880 1.01 6 5 13 4 912 3.12 Melia azedarach 291 53 1 394 54 100 12.00 44 0 82 181 0.98 17 5 29 4 856 4.22 Morus alba 72 6 192 3 986 1.35 2 1 2 11 0.02 6 1 10 481 0.92 Nephrolepis exaltata 10 1 12 318 0.09 2 0 6 131 0.08 9 1 13 337 1.07 Nicotiana glauca 126 7 274 7 812 2.11 51 2 168 3 657 2.33 1 0 1 50 0.09 Opuntia fi cus-indica 330 39 1 159 47 136 10.10 73 5 267 7 242 3.85 8 5 8 4 261 2.48 robusta 50 0 61 191 0.34 12 1 13 1 048 0.26 stricta 82 13 168 5 112 1.32 1 0 1 50 0.02 Paraserianthes lophantha 5 0 10 104 0.06 47 9 274 19 936 5.22 7 0 7 34 0.51 Passifl ora edulis 22 0 40 192 0.23 5 0 6 24 0.07 13 0 19 64 1.37 Pennisetum clandestinum 10 3 12 2 414 0.27 26 4 28 2 106 0.54 3 3 3 3 000 1.56 Pereskia aculeata 34 7 91 5 569 0.96 4 0 4 13 0.05 14 4 39 2 963 4.08 Pinus patula 30 7 90 6 154 1.00 14 5 18 1 194 1.80 pinaster 13 5 34 6 549 0.74 66 36 355 39 368 8.14 9 4 29 4 586 4.10 radiata 3 0 4 53 0.03 53 15 186 8 902 3.08 4 3 20 1 235 1.96 Populus alba 6 0 11 350 0.09 1 0 1 50 0.02 alba/canescens 51 22 171 15 598 2.24 1 0 1 50 0.02 8 2 8 611 0.84 ×canescens 36 8 76 5 390 0.86 82 34 279 24 456 5.74 3 0 4 22 0.29 Prosopis glandulosa 7 5 13 1 421 0.19 2 0 2 51 0.03 Combined taxa e.g. Ageratum conyzoides/houstonianum indicate uncertainty of identifi cation. QSp, quarter-degree squares present; QSa, quarter- degree squares abundant; R, records; A, total weighted abundance (see text); Pv, prominence value (bold numbers: highest prominence values which add up to ± upper 80% of summed values—see text). Bothalia 37,2 (2007) 227

APPENDIX 2.—Prominent invaders in Savanna Biome, Fynbos Biome and Forest habitats (cont.)

Savanna Biome Fynbos Biome Forest Habitats Scientiﬁ c name QSp QSa R A Pv QSp QSa R A Pv QSp QSa R A Pv Prosopis (cont.) glandulosa/velutina 64 5 168 4 869 1.30 22 2 63 1 657 0.90 velutina 6 1 7 257 0.06 3 0 3 52 0.04 Prunus persica 53 0 115 933 0.69 41 0 65 191 0.78 1 0 1 1 0.07 Psidium guajava 124 45 662 53 388 8.07 6 0 9 45 0.11 14 8 25 5 174 4.09 Pyracantha angustifolia 6 0 13 40 0.07 5 0 6 15 0.07 2 0 2 2 0.14 angustifolia/crenulata 5 0 5 5 0.03 2 0 2 2 0.02 1 0 1 50 0.09 Ricinus communis 256 46 1 230 40 996 10.00 87 7 250 4 189 3.35 25 5 30 1 618 2.84 Robinia pseudoacacia 10 0 13 80 0.08 6 0 6 15 0.07 1 0 1 1 0.07 Rosa rubiginosa 8 0 12 128 0.07 4 0 6 162 0.09 2 0 2 51 0.16 Rubus cuneifolius 18 4 45 10 711 1.16 5 3 6 2 451 1.52 fruticosus 15 4 24 2 146 0.31 55 21 188 16 874 3.91 7 3 14 1 701 1.75 pascuus 2 1 2 250 0.03 sp. 31 7 88 8 076 1.16 5 2 5 430 0.10 10 1 12 443 1.04 ×proteus 2 2 2 400 0.05 Salix babylonica 67 5 140 3 609 1.05 38 3 74 1 838 1.05 6 0 6 211 0.52 fragilis 2 0 5 121 0.04 1 1 1 200 0.03 Salsola kali/tragus 22 2 27 1 027 0.23 12 1 13 511 0.20 Schinus molle 52 0 82 463 0.47 24 0 49 358 0.61 1 0 1 10 0.07 Senna didymobotrya 103 25 261 14 239 2.60 3 0 3 3 0.04 12 1 15 406 1.24 Sesbania punicea 139 26 405 21 438 3.97 60 19 175 17 026 3.77 7 0 8 66 0.59 Solanum elaeagnifolium 18 0 18 183 0.11 3 1 3 251 0.06 mauritianum 123 52 748 77 619 10.60 28 2 86 2 035 1.21 66 32 97 27 090 19.00 seaforthianum 28 3 75 2 636 0.62 1 0 1 10 0.01 9 0 31 263 2.30 Tithonia diversifolia 46 5 120 4 033 0.98 8 0 10 313 0.84 Xanthium strumarium 74 16 126 8 577 1.40 2 1 3 251 0.06 4 0 4 62 0.31 Combined taxa e.g. Ageratum conyzoides/houstonianum indicate uncertainty of identiﬁ cation. QSp, quarter-degree squares present; QSa, quarter- degree squares abundant; R, records; A, total weighted abundance (see text); Pv, prominence value (bold numbers: highest prominence values which add up to ± upper 80% of summed values—see text).

APPENDIX 3.—Prominent invaders in Grassland Biome, Nama-Karoo Biome and Succulent Karoo Biome

Grassland Biome Nama-Karoo Biome Succulent Karoo Biome Scientifi c name QSp QSa R A Pv QSp QSa R A Pv QSp QSa R A Pv Acacia cyclops 1 0 1 10 0.04 32 8 105 12 150 26.30 dealbata 206 99 922 116 050 20.90 2 0 2 2 0.06 decurrens 82 24 194 21 677 4.06 longifolia 19 3 31 4 362 0.76 1 0 2 2 0.16 mearnsii 197 116 1 003 114 979 21.30 9 6 36 10 373 18.20 melanoxylon 49 4 109 4 435 1.33 2 0 2 11 0.18 saligna 2 0 2 11 0.02 24 8 72 6 471 15.30 Achyranthes aspera 30 0 34 128 0.26 2 0 2 51 0.09 Agave americana 148 2 219 2 507 1.87 81 3 160 1 961 5.90 17 0 31 255 2.84 sisalana 21 0 25 52 0.19 3 0 3 21 0.10 4 0 4 4 0.32 Ageratum conyzoides 5 0 6 162 0.06 conyzoides/houstonianum 3 2 3 410 0.07 houstonianum 4 0 5 103 0.05 Argemone mexicana 3 0 3 52 0.03 ochroleuca 38 2 56 1 155 0.54 26 4 27 3 549 2.52 2 0 2 20 0.19 sp. 1 0 1 10 0.01 1 0 1 1 0.03 6 0 6 55 0.56 Arundo donax 89 8 188 5 068 1.97 51 8 77 2 927 3.80 28 7 46 5 316 11.50 Atriplex infl ata 3 1 4 460 0.09 83 42 107 37 882 21.00 46 15 61 5 687 13.30 nummularia 4 0 6 153 0.06 88 11 181 4 732 7.90 45 3 83 2 176 9.81 Azolla fi liculoides 88 41 172 21 806 3.92 47 25 70 12 944 8.30 Caesalpinia decapetala 47 19 174 12 902 2.83 Cardiospermum grandifl orum 3 1 3 202 0.05 grandifl orum/halicacabum 2 0 2 11 0.02 Cereus jamacaru 12 0 12 106 0.10 6 0 6 73 0.22 Cestrum laevigatum 16 4 20 3 430 0.57 Combined taxa e.g. Ageratum conyzoides/houstonianum indicate uncertainty of identifi cation. QSp, quarter-degree squares present; QSa, quarter- degree squares abundant; R, records; A, total weighted abundance (see text); Pv, prominence value (bold numbers: highest prominence values which add up to ± upper 80% of summed values—see text). 228 Bothalia 37,2 (2007)

APPENDIX 3.—Prominent invaders in Grassland Biome, Nama-Karoo Biome and Succulent Karoo Biome (cont.)

Grassland Biome Nama-Karoo Biome Succulent Karoo Biome Scientifi c name QSp QSa R A Pv QSp QSa R A Pv QSp QSa R A Pv Chromolaena odorata 14 7 29 4 130 0.72 Cinnamomum camphora 1 0 3 3 0.02 Cirsium vulgare 126 14 253 10 776 3.14 11 2 16 1 463 1.19 Datura ferox 82 9 85 3 429 1.03 21 2 26 2 566 2.03 innoxia 3 2 3 401 0.68 2 0 2 11 0.07 sp. 46 0 65 877 0.57 2 0 2 2 0.06 1 0 1 50 0.15 stramonium 126 12 174 6 224 2.01 40 2 48 1 259 2.10 1 0 1 10 0.09 Eichhornia crassipes 21 21 106 14 000 2.48 Eucalyptus camaldulensis 16 3 17 2 406 0.42 14 2 14 2 195 1.48 27 0 34 347 3.21 diversicolor 2 1 2 210 0.04 2 0 2 2 0.16 grandis 45 8 78 4 638 1.13 sp. 276 12 614 11 153 5.76 21 0 27 188 0.93 14 0 16 102 1.42 Hakea sericea 1 0 1 50 0.01 8 1 11 1 028 2.40 Ipomoea indica 2 0 2 2 0.01 indica/purpurea 15 3 16 774 0.21 purpurea 12 0 13 120 0.11 sp. 1 0 1 1 0.01 Jacaranda mimosifolia 55 0 115 662 0.90 1 0 1 1 0.03 Lantana camara 59 24 207 10 606 2.79 1 0 1 10 0.09 Macfadyena unguis-cati 5 3 5 651 0.12 Melia azedarach 177 12 588 11 198 5.58 36 0 49 250 1.65 3 0 6 6 0.49 Morus alba 54 2 109 1 985 1.02 1 0 1 1 0.03 Nephrolepis exaltata 1 0 1 10 0.01 Nicotiana glauca 48 1 72 1 266 0.67 92 2 206 3 980 8.30 66 2 237 5 417 26.80 Opuntia fi cus-indica 257 10 570 11 437 5.48 161 3 368 5 688 14.00 40 0 81 974 7.87 robusta 72 1 120 927 0.97 80 0 130 1 029 4.60 11 0 13 49 1.10 stricta 12 1 12 229 0.11 9 0 9 18 0.29 2 0 3 3 0.24 Paraserianthes lophantha 2 0 2 2 0.16 Passifl ora edulis 5 0 9 45 0.07 Pennisetum clandestinum 10 4 11 3 354 0.49 2 1 2 1 010 0.54 Pereskia aculeata 6 1 7 206 0.08 Pinus patula 55 6 148 5 482 1.73 pinaster 4 1 4 230 0.06 2 2 8 2 082 3.72 radiata 8 0 10 86 0.08 6 0 6 202 0.78 Populus alba 8 2 10 515 0.13 alba/canescens 128 25 283 18 169 4.26 5 0 5 54 0.18 ×canescens 198 75 486 51 371 9.80 40 8 67 4 537 4.20 15 5 31 1 643 4.89 Prosopis glandulosa 3 0 3 61 0.03 25 4 29 2 395 2.00 3 1 3 1 060 1.81 glandulosa/velutina 29 6 54 4 738 0.97 214 55 666 73 664 56.00 61 10 156 7 823 24.00 velutina 1 0 1 1 0.01 37 5 41 2 788 2.60 1 0 1 10 0.09 Prunus persica 211 1 530 6 232 4.56 13 0 15 33 0.49 1 0 3 12 0.26 Psidium guajava 29 5 60 2 357 0.72 1 0 1 1 0.08 Pyracantha angustifolia 122 3 256 3 661 6.07 9 0 10 19 0.32 angustifolia/crenulata 33 3 44 1 176 0.46 Ricinus communis 80 3 166 3 149 1.57 13 0 24 208 0.85 20 0 31 313 2.92 Robinia pseudoacacia 83 14 145 8 576 2.09 11 0 14 157 0.51 Rosa rubiginosa 104 12 255 11 201 3.20 3 0 3 3 0.10 Rubus cuneifolius 57 31 191 38 602 6.12 fruticosus 15 5 24 3 219 0.57 4 2 8 571 1.48 pascuus 1 1 1 200 0.03 sp. 50 21 86 21 188 3.23 ×proteus 2 1 2 250 0.05 Salix babylonica 310 79 1 069 78 092 17.30 54 2 90 1 555 3.55 6 0 8 26 0.67 fragilis 71 23 169 15 388 3.10 1 0 1 1 0.03 Salsola kali/tragus 20 1 23 610 0.24 95 26 117 12 309 9.50 6 1 7 233 0.90 Schinus molle 54 0 73 360 0.57 75 2 156 3 952 6.80 26 0 47 222 4.06 Senna didymobotrya 33 4 75 2 100 0.79 Sesbania punicea 117 22 238 13 269 3.34 4 0 4 13 0.13 3 1 8 332 1.13 Solanum elaeagnifolium 18 4 21 3 498 0.58 10 6 16 10 202 5.30 2 0 2 2 0.16 mauritianum 114 45 530 55 565 10.60 seaforthianum 1 0 1 10 0.01 Tithonia diversifolia 3 0 3 52 0.03 Xanthium strumarium 64 2 72 2 364 0.81 9 2 11 1 441 1.03 Combined taxa e.g. Ageratum conyzoides/houstonianum indicate uncertainty of identifi cation. QSp, quarter-degree squares present; QSa, quarter- degree squares abundant; R, records; A, total weighted abundance (see text); Pv, prominence value (bold numbers: highest prominence values which add up to ± upper 80% of summed values—see text). Bothalia 37,2 (2007) 229

APPENDIX 4.—Summary of results for all naturalized and casual alien plants in the study area, Savanna Biome, Fynbos Biome, Forest habitats, Grassland Biome, Nama-Karoo Biome, Succulent Karoo Biome and watercourse/wetland habitats

Plant name QSp QSa Study Savanna Fynbos Forest Grassland Nama- Succulent Watercourse/ area Biome Biome habitat Biome Karoo Karoo wetland records records records records records Biome Biome records records records Acacia baileyana 86 135 20 39 1 75 110 cultriformis # 11 1 cyclops 166 91 1 097 181 810 9 1 105 206 dealbata 256 115 1 079 132 23 12 922 2 4 542 decurrens 101 31 232 38 4 194 30 elata 35 1 60 4 51 43 2 4 fi mbriata 11 1 1 implexa 22 2 2 longifolia 94 34 363 66 264 13 31 2 132 mearnsii 428 251 2 620 710 871 91 1 003 36 953 melanoxylon 134 29 482 73 298 85 109 2 130 paradoxa 12 2 podalyriifolia 57 2 101 49 27 1 25 6 pycnantha 35 15 135 4 131 6 saligna 158 93 1 030 96 860 8 2 72 363 viscidula 11 1 Acanthocereus ?tetragonus 11 1 1 Acanthospermum australe * 11 1 hispidum * 111 Acanthus polystachyus var. 111 pseudopubescens # Acer negundo 11 11 1 ? sp. 11 11 1 Achyranthes aspera * 77 3 85 44 512 34 223 Acorus calamus * 11 1 1 Acrocarpus fraxinifolius # 11 11 Adiantum raddianum # 1111 Agave americana var. americana 431 8 761 267 84 2 219 160 31 118 var. expansa 11 1 decipiens # 11 1 sisalana 170 12 293 251 10 4 25 3 4 13 sp. 31 1 60 53 43 12 Ageratina adenophora 11 4 26 11 10 35 10 riparia ?# 111 Ageratum conyzoides 37 8 50 44 16 32 conyzoides/houstonianum 31 8 49 45 1133 24 houstonianum 26 2 31 26 85 7 Agrimonia cf. parvifl ora # 11 1 1 Agrostemma githago * 11 1 Ailanthus altissima 32 2 40 11 6 19 31 9 Albizia chinensis # 111 lebbeck 42 5 5 procera 111 Alhagi maurorum 10 11 81 1 1 3 Alisma plantago-aquatica 81 9 2 7 9 Alnus glutinosa 11 1 1 Alpinia zerumbet 5 7 241 1 2 Alternanthera pungens * 441 3 1 Amaranthus hybridus * 33 3 1 sp.* 111 Ambrosia artemisiifolia * 221 1 1 Ammi majus * 11 1 Anigozanthos fl avidus # 11 1 1 Anredera cordifolia 24 2 25 17 37 5 4 Antigonon leptopus 355 1 Apium graveolens * 111

Combined taxa e.g. Ageratum conyzoides/houstonianum indicate uncertainty of identiﬁ cation. #, casual alien plants: occurring outside cultivation; some species ﬂ ourishing but less than 10 years of records in SAPIA precludes being categorized as ‘naturalized’ (Pyšek et al. 2004). *, mainly herbaceous species that are suspected of being under-estimated in this survey. QDSp, quarter-degree squares present; QDSa, quarter-degree squares abundant. Bold numbers in biome categories add up to upper 80% or more of total records. 230 Bothalia 37,2 (2007)

Plant name QSp QSa Study Savanna Fynbos Forest Grassland Nama- Succulent Watercourse/ area Biome Biome habitat Biome Karoo Karoo wetland records records records records records Biome Biome records records records Araujia sericifera 36 1 53 15 24 35 112 Ardisia crenata 21 3 3 2 3 Argemone mexicana 27 5 36 32 11 3 20 ochroleuca subsp. ochroleuca 154 20 206 119 21 56 27 2 50 sp. 14 2 15 7 11 6 9 Aristolochia elegans 61 8 7 2 1 4 Arundo donax 371 82 855 372 172 3 188 77 46 548 Astartea fascicularis # 11 1 Atriplex infl ata 164 77 213 16 25 4 107 61 16 muelleri * 11 1 nummularia subsp. nummularia 172 18 333 18 45 6 181 83 107 semibaccata * 45 321 sp. 10 10 8 2 2 Azolla fi liculoides 194 92 354 75 37 2 172 70 354 ?pinnata var. imbricata 31 6 6 4 sp. 4 8 8 8 Baeckia sp. # 11 1 Bambusa balcooa 32 50 42 51 3 43 sp. # 1 1 1 Bambuseae sp. 8 9 7 1 1 3 Banksia ericifolia # 11 1 integrifolia # 11 1 1 Bauhinia purpurea 111 sp. 1 2 2 variegata 91111 1 Begonia cucullata # 1211 Bidens bipinnata * 23 23 23 7 biternata * 111 pilosa * 39 3 65 17 1 47 12 Billardiera heterophylla # 11 1 Boerhavia erecta * 11 1 Briza maxima * 111 Bromus catharticus * 22 111 diandrus * 11 1 1 pectinatus * 11 1 1 Brugmansia ×candida 6 7 161 Bryophyllum delagoense 41 6 6 ?Buddleja madagascariensis # 11 1 Caesalpinia decapetala 127 41 413 239 25 174 153 gilliesii 18 19 63822 Callistemon citrinus # 11 1 glaucus # 111 rigidus 111 sp. 1 1 1 1 viminalis 11 1 1 Calotropis procera # 111 Campuloclinium macrocephalum 14 5 25 16 1 9 3 Canna glauca # 1111 indica 26 1 34 19 7 6 8 11 sp. 13 17 10 4 1 3 10 ×generalis 7881 5 Capsella bursa-pastoris * 11 1 Cardiospermum grandifl orum 43 12 57 54 73 29 Combined taxa e.g. Ageratum conyzoides/houstonianum indicate uncertainty of identifi cation. #, casual alien plants: occurring outside cultivation; some species fl ourishing but less than 10 years of records in SAPIA precludes being categorized as ‘naturalized’ (Pyšek et al. 2004). *, mainly herbaceous species that are suspected of being under-estimated in this survey. QDSp, quarter-degree squares present; QDSa, quarter-degree squares abundant. Bold numbers in biome categories add up to upper 80% or more of total records. Bothalia 37,2 (2007) 231

Plant name QSp QSa Study Savanna Fynbos Forest Grassland Nama- Succulent Watercourse/ area Biome Biome habitat Biome Karoo Karoo wetland records records records records records Biome Biome records records records Cardiospermum (cont.) grandiﬂ orum/halicacabum 16 5 20 18 42 5 halicacabum 27 35 34 51 26 Carica papaya 666 2 Castanea dentata ?# 11 11 Castanospermum australe ?# 111 Casuarina cunninghamiana 91282 2 8 cunninghamiana/equisetifolia 42 2 63 46 61 11 23 equisetifolia 24 1 57 55 62 7 Catharanthus roseus 38 1 53 49 12 3 11 Cedrus deodara ?# 55 5 Celtis australis † ?? ? occidentalis † ?? ? sinensis † 11 1 ? Cenchrus brownii * 111 Centranthus ruber ?# 22 2 2 Cereus jamacaru 124 11 193 169 61 126 6 Cestrum aurantiacum 81 10 7 16 2 2 aurantiacum/laevigatum 73 8 5 4 3 1 elegans 22 2 2 2 laevigatum 70 16 167 138 930 20 41 parqui 354 1 sp. 1 1 1 1 Chamaesyce prostrata * 331 2 serpens * 11 1 Chenopodium album * 33 3 Chorizema cordatum # 11 1 Chromolaena odorata 93 64 558 529 77 29 220 Cichorium intybus 12 1 16 14 22 Cinnamomum camphora 10 1 19 13 3 14 3 3 Cirsium arvense * 22 2 vulgare 188 20 345 68 8 253 16 40 Citrus limon 111 sp. 5 5 1 1 4 2 Coix lacryma-jobi * 1211 Colocasia esculenta 11 3 19 14 5 119 Commelina benghalensis * 915735 3 21 Convolvulus arvensis 23 1 23 44 114 2 Conyza bonariensis * 441 3 1 canadensis * 332 1 1 primulifolia * 111 sp.* 332 1 2 sumatrensis * 11 1 Coreopsis lanceolata 11 15 11 4 Cortaderia jubata 77 2 5 1 jubata/selloana 23 2 28 11 1 16 14 selloana 54 1 104 15 77 5 10 11 21 Corymbia ﬁ cifolia ?# 3312 Cosmos bipinnatus * 48 10 122 3 119 2 Cotoneaster coriaceus ?# 11 1 franchetii 71 7 2 1 5 1 glaucophyllus 221 1 franchetii/pannosus 19 27 1 2 1 23 13 pannosus 25 30 2 1 25 23

Combined taxa e.g. Ageratum conyzoides/houstonianum indicate uncertainty of identiﬁ cation. #, casual alien plants: occurring outside cultivation; some species ﬂ ourishing but less than 10 years of records in SAPIA precludes being categorized as ‘naturalized’ (Pyšek et al. 2004). *, mainly herbaceous species that are suspected of being under-estimated in this survey. †, Celtis australis (probably naturalized), C. occidentalis (probably naturalized) and C. sinensis (naturalized) easily mistaken for indigenous C. africana and suspected of being under-estimated in this survey. QDSp, quarter-degree squares present; QDSa, quarter-degree squares abundant. Bold numbers in biome categories add up to upper 80% or more of total records. 232 Bothalia 37,2 (2007)

Plant name QSp QSa Study Savanna Fynbos Forest Grassland Nama- Succulent Watercourse/ area Biome Biome habitat Biome Karoo Karoo wetland records records records records records Biome Biome records records records Cotoneaster (cont.) sp. 21 23 1 1 21 4 Crataegus sp. # 2 2 2 ×lavallei 44 14 Crotalaria agatifl ora subsp. 18 29 24 81 agatifl ora Cryptomeria japonica # 111 1 Cryptostegia grandifl ora 133 3 Cuphea ignea # 11 1 Cupressus arizonica 47 68 1 62 51 lusitanica 221 1 1 sp. 18 24 3 11 19 12 Cuscuta campestris 82 1 103 22 32 73 523 campestris/suaveolens 34 40 22 6 7 66 5 suaveolens 7722411 Cydonia oblonga 77 3 311 Cytisus scoparius 10 15 1 14 1 Dahlia imperialis ?# 122 Datura ferox 175 14 201 87 32 85 26 55 innoxia 29 3 36 30 11 3 2 18 sp. 84 1 110 41 1 65 21 7 stramonium 286 24 373 131 19 6 174 48 1 73 Delonix regia 5552 Desmanthus virgatus * 111 Dracocephalum canariense 11 1 Duranta erecta 32 35 33 82 8 Dysphania ambrosioides * 221 1 1 Echinopsis spachiana 57 2 83 29 2 14 37 1 Echium plantagineum 51 6 64 11 31 19 34 plantagineum/vulgare 19 2 17 1 vulgare 29 2 31 1 921 2 Egeria densa 222 1 Eichhornia crassipes 87 72 431 279 46 106 431 Eragrostis pilosa * 111 Eriobotrya japonica 3 3 121 1 Eucalyptus camaldulensis 121 22 182 36 81 3171434 137 cinerea 11 13 2 11 cladocalyx 37 4 83 2 71 10 26 cloeziana 111 conferruminata 41 10 117 10 106 112 diversicolor 49 8 153 1 148 28 2 2 32 ?exserta 11 1 1 1 fastigata 111 globulus 12 16 16 51 gomphocephala 611 11 grandis 100 16 190 111 119 78 67 leucoxylon ?# 22 2 2 microcorys 12 2 microtheca 11 1 1 paniculata 111 regnans 67 7 3 robusta ?# 111 1 sideroxylon ?# 11 1 sp. 505 30 1 103 299 147 17 614 27 16 266 tereticornis ?# 11 1 1 Eugenia unifl ora 222 Euphorbia heterophylla 332 1 2 peplus * 11 1 pulcherrima ?# 699 1 Combined taxa e.g. Ageratum conyzoides/houstonianum indicate uncertainty of identifi cation. #, casual alien plants: occurring outside cultivation; some species fl ourishing but less than 10 years of records in SAPIA precludes being categorized as ‘naturalized’ (Pyšek et al. 2004). *, mainly herbaceous species that are suspected of being under-estimated in this survey. QDSp, quarter-degree squares present; QDSa, quarter-degree squares abundant. Bold numbers in biome categories add up to upper 80% or more of total records. Bothalia 37,2 (2007) 233

Plant name QSp QSa Study Savanna Fynbos Forest Grassland Nama- Succulent Watercourse/ area Biome Biome habitat Biome Karoo Karoo wetland records records records records records Biome Biome records records records Fallopia convolvulus * 11 1 Ficus carica 18 26 1 19 21 3 13 elastica # 111 macrophylla # 111 pumila 221 1 Flaveria bidentis * 12 12 11 11 6 Foeniculum vulgare * 91035 11 1 Fraxinus americana 13 15 11 4 3 angustifolia 3311 1 sp. 13 14 11 3 4 Fuchsia sp. # 11 11 Genista monspessulana 22 2 Glandularia aristigera * 14 27 1 26 1 ×hybrida ?# 11 1 Glebionis coronaria 21 4 2 2 Gleditsia triacanthos 111 1 162 12 1 136 12 1 41 Gnaphalium luteoalbum * 111 1 Gomphrena celosioides * 22 2 1 Grevillea robusta 53 80 68 2 9 10 22 rosmarinifolia # 11 1 sericea # 11 1 Guilleminea densa * 22 2 Hakea drupacea 28 2 58 4 53 1 gibbosa 18 3 34 2 31 1 salicifolia 51 5 2 3 1 sericea 77 17 230 14 204 31 115 victoriae # 11 1 Harrisia martinii 21 10 33 32 13 Hedychium coccineum 31 6 6 2 2 coronarium 14 1 19 18 21 2 ﬂ avescens 525 321 2 gardnerianum 12 18 11 4 7 1 sp. 7 2 8 4 3 2 1 4 Helianthus annuus * 588 Heliotropium amplexicaule * 232 1 Hibiscus trionum * 22 2 Homalanthus populifolius 22 22 Hordeum murinum * 11 1 1 Hylocereus undatus 8871 Hypericum patulum 11 2 1 2 1 perforatum 13 1 19 16 33 Hypochaeris radicata * 11 1 Ipomoea alba 22 1 32 28 54 19 carnea subsp. ﬁ stulosa 21 1 36 35 17 indica 23 3 27 22 33 2 8 indica/purpurea 74 7 120 86 18 16 16 34 nil 11 1 1 purpurea 37 3 46 23 10 9 13 12 sp. 3 3 2 1 1 Jacaranda mimosifolia 195 16 613 497 21 115 1 105 Jasminum humile 23 3 mesnyi ?# 11 1 Jatropha gossypiifolia 41010 1 sp. 221 1

Plant name QSp QSa Study Savanna Fynbos Forest Grassland Nama- Succulent Watercourse/ area Biome Biome habitat Biome Karoo Karoo wetland records records records records records Biome Biome records records records Juniperus pinchotii # 11 1 sp. 4 5 5 3 virginiana 17 2 28 1 27 8 Lactuca serriola * 11 1 Lagerstroemia indica 676 1 Lantana camara 247 116 2 111 1 843 60 63 207 1 289 Lemna gibba 32 3 2 1 3 sp. 3 1 3 1 1 1 3 Lepidium didymum * 111 draba 44 13 Leptospermum laevigatum 38 15 102 6 96 310 Leucaena leucocephala 36 3 123 115 11 8 43 Ligustrum japonicum 78226 4 lucidum 12 1 16 5 2 11 6 ovalifolium 31 3 2 1 1 1 sinense 8111 10 5 sp. 12 2 13 6 17 7 vulgare 31 5 3 2 5 Lilium formosanum 15 3 30 18 12 Limonium sinuatum 10 1 10 5412 Linaria genistifolia * 11 1 maroccana * 13 3 Litsea glutinosa 83 10 10 65 Lonicera japonica var. halliana 55213 4 Lygodium japonicum # 1111 Lythrum salicaria 11 1 1 Macfadyena unguis-cati 22 9 52 47 13 5 14 Maireana brevifolia ?# 111 Malus pumila var. paradisi- 57 2 5 1 aca ?# Malva dendromorpha 16 1 24 1 19 1 3 3 linnaei * 11 1 parviﬂ ora * 22 1 1 Malvastrum coromandelianum * 221 1 Mangifera indica 12 1 30 30 116 Manihot esculenta 8109 11 grahamii # 6771 1 Medicago sativa * 221 1 Melaleuca hypericifolia 12 2 wilsonii # 11 1 Melia azedarach 551 65 2 119 1 394 82 29 588 49 6 674 Melilotus alba * 15 26 2 24 2 Metasequioa glyptostroboides # 11 1 1 Metrosideros excelsa 21 5 5 3 Mimosa pigra 61 8 7 1 7 pudica var. hispida * 222 Mirabilis jalapa * 773 4 Momordica charantia * 111 Monstera deliciosa # 111 1 Montanoa hibiscifolia 24 2 46 41 65 13 Moringa oleifera # 222 Morus alba 129 8 304 192 210 109 1 164 Musa sp. # 81515 10 Myoporum tenuifolium subsp. 30 49 2 44 134 montanum

Plant name QSp QSa Study Savanna Fynbos Forest Grassland Nama- Succulent Watercourse/ area Biome Biome habitat Biome Karoo Karoo wetland records records records records records Biome Biome records records records Myriophyllum aquaticum 48 10 81 39 17 25 81 spicatum 20 23 11 2 9 123 Nassella tenuissima 12 2 1 trichotoma 12 3 16 12 1 10 11 Nasturtium offi cinale 50 1 64 19 41 38 364 Nephrolepis exaltata 13 1 19 12 6 13 1 2 Nerium oleander 23 2 46 16 21 14 4 36 Nicandra physalodes * 111 Nicotiana glauca 383 14 957 274 168 172206 237 441 tabacum ?# 333 3 Nymphaea ×marliacea 11 1 1 mexicana 22 4 1 3 4 Oenothera biennis * 19 1 19 513110 glazioviana * 11 1 1 indecora * 11 1 jamesii * 15 2 17 7 1 9 10 laciniata * 11 1 rosea * 44 4 1 sp. 4 4 4 tetraptera * 11 1 Olyra latifolia * 1111 Opuntia aurantiaca 61 3 84 50 28 42 4 engelmannii (= O. lindheimeri) 10 3 15 6 2 7 1 exaltata 662 3 fi cus-indica 861 57 2 445 1 159 267 8 570 368 81 129 fulgida 11 2 12 7 2 3 humifusa 25 3 32 18 2 10 22 ?humifusa/engelmannii 48 4 49 23 17 91 imbricata 131 15 151 49 83 48 43 36 microdasys 91043 3 monacantha 48 1 114 90 22 52 17 robusta 225 2 337 61 13 120 130 13 4 sp. 37 39 21 5 10 32 spinulifera ?# 1 1 1 stricta 106 14 193 168 112934 ?stricta × humifusa 111 Orobanche minor 4523 Oxalis corniculata * 441 3 1 Pandanus sp. # 111 1 Paraserianthes lophantha 54 9 286 10 274 7282 Parkinsonia aculeata 15 18 12 6 4 Parthenium hysterophorus 15 3 29 29 8 Parthenocissus quinquefolia # 111 Paspalum dilatatum * 6 6 122 3 1 quadrifarium * 11 1 1 urvillei * 111 Passifl ora caerulea 12 20 10 9 21 12 edulis 32 55 40 619 9 12 sp. 19 1 22 13 6 11 3 7 suberosa 67621 subpeltata 21 1 26 22 34 3 tripartita var. mollissima 44 311 1 Pennisetum clandestinum 48 12 56 12 28 3 14 26 purpureum 40 6 87 82 530 setaceum 66 17 84 39 20 8 17 2 setaceum/villosum 15 1 15 44 6 11 sp. 11 4 15 11 villosum 22 5 26 2 3 14 7 1 Combined taxa e.g. Ageratum conyzoides/houstonianum indicate uncertainty of identifi cation. #, casual alien plants: occurring outside cultivation; some species fl ourishing but less than 10 years of records in SAPIA precludes being categorized as ‘naturalized’ (Pyšek et al. 2004). *, mainly herbaceous species that are suspected of being under-estimated in this survey. QDSp, quarter-degree squares present; QDSa, quarter-degree squares abundant. Bold numbers in biome categories add up to upper 80% or more of total records. 236 Bothalia 37,2 (2007)

Plant name QSp QSa Study Savanna Fynbos Forest Grassland Nama- Succulent Watercourse/ area Biome Biome habitat Biome Karoo Karoo wetland records records records records records Biome Biome records records records Pereskia aculeata 44 8 102 91 4397 8 Persea americana # 22 2 1 Persicaria lapathifolia * 11 1 1 Phoenix canariensis 33 2 13 dactylifera 44 3 1 4 Phormium tenax # 22 2 2 Physalis peruviana * 332 1 viscosa * 331 2 2 Phytolacca dioica 30 38 17 20 1115 icosandra * 45 5 1 Pinus canariensis 61 9 2 7 elliottii 34 6 59 35 3 23 17 elliottii/taeda 30 1 59 41 2 18 9 halepensis 85 3 136 45 53 33 41 9 patula 85 13 238 90 18 148 59 pinaster 85 44 401 34 355 29 4 8 36 pinea 18 35 1 30 41 radiata 70 15 206 4 186 20 10 6 16 roxburghii 22 2 sp. 126 14 169 47 47 3 70 32 18 taeda 71165 3 Pistia stratiotes 24 6 63 60 21 63 Pittosporum undulatum 37 71 3 Pityrogramma calomelanos * 1111 Plantago lanceolata * 44 4 1 major * 221 1 virginica * 11 1 Platanus sp. # 11 1 Plectranthus comosus 19 22 811 1 3 2 Polygonum aviculare * 11 1 1 Polypogon monspeliensis * 111 Pomaderris kumeraho # 11 1 Pontederia cordata 222 2 Populus alba 15 2 22 11 1 10 17 alba/×canescens 185 47 460 171 18283 5 336 deltoides 100 6 169 37 31 114 15 117 nigra var. italica 90 120 2 7 94 17 100 ×canescens 371 130 939 76 279 4 486 67 31 823 Portulaca oleracea * 221 1 Prosopis glandulosa var. torreyana 40 10 50 13 2329 313 glandulosa/velutina 390 78 1 108 168 63 54 666 156 443 velutina 48 6 53 7 3141 118 Prunus armeniaca 32 44 1 63421 9 persica 319 1 728 115 65 1 530 15 3 148 serotina 11 1 1 Psidium cattleianum 58741 1 guajava 160 50 732 662 9 25 60 1 179 guineense 2221 sp. 36 7 47 44 63 12 ×durbanensis 2332 Pterocarya stenoptera # 1111 1 Pueraria montana var. lobata 332 1 2 Punica granatum 8111433 2 Pyracantha angustifolia 142 3 285 13 6 2 256 10 78 angustifolia/crenulata 40351521 44 5

Plant name QSp QSa Study Savanna Fynbos Forest Grassland Nama- Succulent Watercourse/ area Biome Biome habitat Biome Karoo Karoo wetland records records records records records Biome Biome records records records Pyracantha (cont.) coccinea 67 7 2 crenulata 22 31 2 1 29 2 Pyrus sp. ?# 55 4 1 1 Quercus canariensis ?# 11 1 cerris ?# 22 2 palustris 55 2 3 1 robur 50 2 88 3 57 5 26 253 sp. 4 6 1 4 1 3 suber ?# 11 1 Richardia brasiliensis * 111 1 humistrata * 11 1 Ricinus communis 456 56 1 701 1 230 250 30 166 24 31 582 Rivina humilis 7 1 10 10 4 1 Robinia pseudoacacia 110 14 178 13 6 1 145 14 66 Rosa multifl ora 553 2 1 ? ×odorata # 111 1 rubiginosa 119 12 276 12 6 2 255 359 sp. # 3 3 2 1 Rubus cuneifolius 75 35 236 45 6 191 71 fl agellaris 34 4 1 fruticosus 89 32 244 24 188 14 24 871 ?pascuus 32 3 2 1 phoenicolasius 44 14 rosifolius 14 1 14 9 23 3 3 sp. 86 30 179 88 512 86 54 ×proteus 43 4 2 2 3 Rumex acetosella subsp. pyrenaicus * 11 1 1 crispus * 221 1 usambarensis 41 4 3 1 Saccharum offi cinarum 15 1 26 25 1 8 Salix babylonica 475 89 1 381 140 74 6 1 069 90 8 1 323 caprea 912 1210 fragilis 75 24 176 5 1 169 1 175 Salsola kali/tragus 155 31 187 27 13 23 117 72 Salvinia molesta 29 7 44 25 14 544 Sambucus canadensis 31 3 1 2 3 sp. 10 11 2 9 6 Scheffl era actinophylla # 111 Schinus molle 231 2 407 82 49 1 73 156 47 136 terebinthifolius 30 2 90 85 29 3 54 Schizolobium parahyba var. 111 parahyba # Schkuhria pinnata * 441 3 1 Senna bicapsularis 16 1 45 45 31 corymbosa 442 2 1 didymobotrya 139 29 339 261 315 75 115 hirsuta 91010 1 multiglandulosa 11 1 12 6412 3 obtusifolia 455 3 occidentalis 56 4 75 74 11 27 pendula var. glabrata 19 2 21 19 22 6 septemtrionalis 63 102 84 1 8 17 31 sp. 16 23 12 6 2 5 12 Sesbania bispinosa * 11 1 1 punicea 323 68 830 405 175 8 238 4 8 500 Combined taxa e.g. Ageratum conyzoides/houstonianum indicate uncertainty of identifi cation. #, casual alien plants: occurring outside cultivation; some species fl ourishing but less than 10 years of records in SAPIA precludes being categorized as ‘naturalized’ (Pyšek et al. 2004). *, mainly herbaceous species that are suspected of being under-estimated in this survey. QDSp, quarter-degree squares present; QDSa, quarter-degree squares abundant. Bold numbers in biome categories add up to upper 80% or more of total records. 238 Bothalia 37,2 (2007)

Plant name QSp QSa Study Savanna Fynbos Forest Grassland Nama- Succulent Watercourse/ area Biome Biome habitat Biome Karoo Karoo wetland records records records records records Biome Biome records records records Sigesbeckia orientalis * 111 Silybum marianum * 797 2 Sisymbrium orientale * 11 1 Sisyrinchium sp. * 22 22 Solanum betaceum 45 431 capsicoides * 111 chrysotrichum 21 21 21 12 elaeagnifolium 51 11 60 18 3 21 16 24 mauritianum 265 99 1 364 748 86 97 530 419 pseudocapsicum * 61018 71 5 seaforthianum 30 3 77 75 1311 50 sisymbriifolium 40 3 51 18 31 30 5 sp. 4 6 5 2 1 1 torvum * 111 Sonchus oleraceus * 331 2 Sophora cf. davidii # 11 1 1 Sorghum halepense 41 2 46 26 32 11 617 Spartium junceum 20 3 43 4 35 312 Spathodea campanulata 344 Sphagneticola trilobata * 122 1 Spiraea cantoniensis # 11 1 Stellaria media * 11 1 Stenocarpus sinuatus ?# 11 1 1 Styphnolobium japonicum ?# 11 1 1 Symphyotrichum squamatum * 11 1 Syncarpia glomulifera 222 Syzygium cumini 91 14 13 21 4 jambos 3321 2 paniculatum 333 Tabebuia chrysotricha ?# 111 Tagetes minuta * 47 78 11 1 66 116 Tamarix chinensis 44 132 ramosissima 7813 4 8 sp. 85 4 110 10 16 8 58 18 85 Taraxacum ofﬁ cinale * 11 1 Tecoma stans 57 4 99 95 24 17 Tephrocactus ?aoracanthus 11 1 1 articulatus 11 1 sp. 1 1 1 Thevetia peruviana 15 23 23 6 Tipuana tipu 24 1 42 33 9 3 Tithonia diversifolia 49 5 123 120 10 3 33 diversifolia/rotundifolia 31 3 3 rotundifolia 19 5 28 22 2 6 7 Toona ciliata 23 1 54 51 73 10 Torilis arvensis * 111 Toxicodendron succedaneum 12 1 14 12 32 3 Tragopogon dubius * 11 1 Tridax procumbens * 111 1 Triplaris americana ?# 111 Triticum aestivum * 11 1 Tropaeolum majus * 3 3 211 1 Ulex europaeus 91421113 Ulmus parvifolia 22 2 1 procera # 11 1 1 sp. # 4 4 3 1 1 Verbena bonariensis * 58 4 115 22 93 13 brasiliensis/bonariensis 243 1 brasiliensis * 222

Plant name QSp QSa Study Savanna Fynbos Forest Grassland Nama- Succulent Watercourse/ area Biome Biome habitat Biome Karoo Karoo wetland records records records records records Biome Biome records records records Verbena (cont.) offi cinalis * 22 2 1 rigida var. rigida * 111 Verbesina encelioides * 18 21 14 7 5 Vinca major * 11 1 Vitis sp. # 612 11 1 Washingtonia sp. ?# 332 13 Wigandia urens var. 3422 caracasana # Wisteria fl oribunda # 11 1 Xanthium sp. 6 7 5 2 1 7 spinosum 83 6 104 30 3 62 827 strumarium 149 21 212 126 34 72 11 95 Yucca aloifolia 20 22 82 9 2 5 Zinnia peruviana * 46511 Combined taxa e.g. Ageratum conyzoides/houstonianum indicate uncertainty of identifi cation. #, casual alien plants: occurring outside cultivation; some species fl ourishing but less than 10 years of records in SAPIA precludes being categorized as ‘naturalized’ (Pyšek et al. 2004). *, mainly herbaceous species that are suspected of being under-estimated in this survey. QDSp, quarter-degree squares present; QDSa, quarter-degree squares abundant. Bold numbers in biome categories add up to upper 80% or more of total records.

APPENDIX 5.—Species checklist The following 601naturalized and casual alien (#) plant species were catalogued in the SAPIA database up to May 2006. Accepted names in roman type. Synonyms in italics. *, taxa added to SAPIA after 2000; †, taxa recorded only in Zimbabwe and Malawi. PRE, species records from the Pretoria National Herbarium Acacia var. expansa (Jacobi) Gentry (= A. expansa Jacobi), Agavaceae, baileyana F.Muell., Fabaceae, Bailey’s wattle spreading century plant cultriformis A.Cunn. ex G.Don, Fabaceae, knife-leaved wattle # decipiens Baker (= A. laxifolia Baker), Agavaceae, false sisal # cyclops A.Cunn. ex G.Don, Fabaceae, red eye sisalana Perrine, Agavaceae, sisal dealbata Link., Fabaceae, silver wattle sp., Agavaceae decurrens Willd., Fabaceae, green wattle elata A.Cunn. ex Benth. (A. terminalis (Salisb.) J.F.Macb. misapplied Ageratina in South Africa), Fabaceae, peppertree wattle adenophora (Spreng.) R.M.King & H.Rob. (= Eupatorium adenophorum fi mbriata A.Cunn. ex G.Don, Fabaceae, fringed wattle Spreng.), Asteraceae, crofton weed implexa Benth., Fabaceae, hickory wattle riparia (Regel) R.M.King & H.Rob. (= Eupatorium riparium Regel), longifolia (Andrews) Willd., Fabaceae, long-leaved wattle Asteraceae, creeping crofton weed ?# mearnsii De Wild., Fabaceae, black wattle Ageratum melanoxylon R.Br., Fabaceae, Australian blackwood conyzoides L., Asteraceae, invading ageratum paradoxa DC. (= A. armata R.Br.), Fabaceae, kangaroo thorn houstonianum Mill., Asteraceae, Mexican ageratum podalyriifolia A.Cunn. ex G.Don, Fabaceae, pearl acacia pycnantha Benth., Fabaceae, golden wattle Agrimonia cf. parvifl ora Aiton, Rosaceae, agrimony # saligna (Labill.) H.L.Wendl. (= A. cyanophylla Lindl.), Fabaceae, Port Agrostemma githago L., Caryophyllaceae, corn cockle Jackson willow Ailanthus altissima (Mill.) Swingle, Simaroubaceae, tree-of-heaven stricta (Andrews) Willd., Fabaceae, hop wattle ?#, *2004 Albizia viscidula Benth., Fabaceae, sticky wattle chinensis (Osbeck) Merr. (= A. stipulata (DC.) Boivin), Fabaceae, Acanthocereus ?tetragonus (L.) Hummelinck, Cactaceae, barbed-wire Chinese false-thorn # cactus lebbeck (L.) Benth., Fabaceae, lebbeck tree procera (Roxb.) Benth., Fabaceae, false lebbeck Acanthospermum australe (Loefl .) Kuntze (= A. brasilum Schrank), Asteraceae, eight- Alhagi maurorum Medik. (= A. camelorum Fisch.), Fabaceae, camelthorn seeded prostrate starbur bush hispidum DC., Asteraceae, upright starbur Alisma plantago-aquatica L., Alismataceae, water plantain Alnus glutinosa (L.) Gaertn. (= A. barbata C.A.Mey.), Betulaceae, Acanthus polystachyus Delile var. pseudopubescens Cufod. (= A. black elder pubescens Engl.), Acanthaceae, bear’s breeches # Alpinia zerumbet (Pers.) B.L.Burtt & R.M.Sm. (= A. speciosa (J.C.Wendl.) Acer K.Schum.), Zingiberaceae, shell ginger buergerianum Miq., Aceraceae, Chinese maple, *2003 # Alternanthera pungens Kunth (= A. repens (L.) Link.), Amaranthaceae, negundo L. (= A. californicum D.Dietr.), Aceraceae, ash-leaved maple khaki bur weed ? sp., Aceraceae, ?red-leafed maple Amaranthus Achyranthes aspera L. (= A. argentea Lam.), Amaranthaceae, burweed hybridus L. , Amaranthaceae, pigweed Acorus calamus L., Acoraceae, calamus sp., Amaranthaceae Acrocarpus fraxinifolius Wight ex Arn., Fabaceae, shingle tree # Ambrosia artemisiifolia L., Asteraceae, annual ragweed Adiantum raddianum C.Presl, Adiantaceae, maidenhair fern # Ammi majus L. (= A. glaucifolium L.), Apiaceae, bishop’s weed Agave Anigozanthos fl avidus DC., Haemodoraceae, yellow kangaroo paw # americana L Anredera cordifolia (Ten.) Steenis, (A. baselloides (Kunth) Baill. var. americana, Agavaceae, American agave misapplied in South Africa), Basellaceae, bridal wreath 240 Bothalia 37,2 (2007)

APPENDIX 5.—Species checklist (cont.)

Antigonon leptopus Hook. & Arn., Polygonaceae, coral creeper Caesalpinia Apium graveolens L., Apiaceae, wild celery decapetala (Roth) Alston (= C. sepiaria Roxb.), Fabaceae, Mauritius thorn Araucaria gilliesii (Hook.) D.Dietr., Fabaceae, bird-of-paradise sp., Araucariaceae, monkey puzzle tree, *2003 # pulcherrima (L.) Sw., Fabaceae, pride of barbados, *2004 # bidwillii Hook., Araucariaceae, bunya-bunya, *2005 # Araujia sericifera Brot., Asclepiadaceae, moth catcher Callisia repens (Jacq.) L., Commelinaceae, creeping inch plant, *2006 # Ardisia Callistemon crenata Sims (A. crispa (Thunb.) A.DC. misapplied in South Africa), citrinus (Curtis) Skeels, Myrtaceae, crimson bottlebrush # Myrsinaceae, coralberry tree glaucus (Curtis) Sweet (= C. speciosus auct.), Myrtaceae, Albany elliptica Thunb. (= A. humilis Vahl), Myrsinaceae, shoebutton ardisia, bottlebrush # *2005 # rigidus R.Br., Myrtaceae, stiff bottlebrush sp., Myrtaceae Argemone viminalis (Sol. ex Gaertn.) G.Don, Myrtaceae, weeping bottlebrush mexicana L., Papaveraceae, yellow-fl owered Mexican poppy ochroleuca Sweet subsp. ochroleuca, Papaveraceae, white-fl owered Calotropis procera (Aiton) W.T.Aiton (= Asclepias procera Aiton), Ascle- Mexican poppy piadaceae, madar # sp., Papaveraceae Campuloclinium macrocephalum (Less.) DC. (= Eupatorium macrocephalum Less.), Asteraceae, pom pom weed Aristolochia elegans Mast., Aristolochiaceae, calico fl ower Arundo donax L., Poaceae, giant reed Canna Astartea fascicularis (Labill.) DC., Myrtaceae # glauca L., Cannaceae, yellow-fl owered glaucous canna # indica L. (= C. edulis Ker Gawl.), Cannaceae, Indian canna Atriplex sp., Cannaceae infl ata F.Muell. (= A. lindleyi Moq. subsp. infl ata (F.Muell.) Paul G.Wilson), ×generalis L.H.Bailey, Cannaceae, garden canna Chenopodiaceae, sponge-fruit saltbush muelleri Benth., Chenopodiaceae, Mueller’s saltbush Capsella bursa-pastoris (L.) Medik., Brassicaceae, shepherd’s purse nummularia Lindl. subsp. nummularia, Chenopodiaceae, old-man Cardiospermum saltbush grandifl orum Sw., Sapindaceae, balloon vine semibaccata R.Br., Chenopodiaceae, Australian saltbush halicacabum L., Sapindaceae, heart pea sp., Chenopodiaceae Carica papaya L. (= Papaya carica Gaertn.), Caricaceae, pawpaw Azolla Castanea dentata (Marshall) Borkh., Fagaceae, American chestnut ?# fi liculoides Lam., Azollaceae, red water fern Castanospermum australe A.Cunn. & C.Fraser ex Hook., Fabaceae, ?pinnata R.Br. subsp. asiatica R.M.K Saunders & K.Fowler (= A. Australian chestnut ?naturalized imbricata (Roxb. ex Griff.) Nakai), Azollaceae, mosquito fern sp., Azollaceae Casuarina cunninghamiana Miq., Casuarinaceae, beefwood Baeckia sp., Myrtaceae # equisetifolia L., Casuarinaceae, horsetail tree Bambusa Catharanthus roseus (L.) G.Don (= Lochnera rosea (L.) Rchb., Vinca balcooa Roxb., Poaceae, common bamboo rosea L.), Apocynaceae, Madagascar periwinkle sp. with tall yellow stems and green leaves, Poaceae, bamboo Cedrus deodara (Roxb. ex D.Don) G.Don, Pinaceae, deodar ?# Bambuseae sp., Poaceae, bamboo Celtis australis L., Ulmaceae, European hackberry ?naturalized Banksia occidentalis L., Ulmaceae, common hackberry ?naturalized ericifolia L.f., Proteaceae, heath banksia # sinensis Pers., Ulmaceae, Chinese nettle tree integrifolia L.f., Proteaceae, coast banksia # Cenchrus brownii Roem. & Schult. (= C. viridis Spreng.), Poaceae, fi ne Bauhinia burgrass purpurea L., Fabaceae, butterfl y orchid tree Centranthus ruber (L.) DC., Valerianaceae, red valerian ?naturalized sp., Fabaceae Cereus jamacaru DC. (C. peruvianus (L.) Mill. misapplied in South variegata L., Fabaceae, orchid tree Africa), Cactaceae, queen of the night Begonia cucullata Willd. (= B. semperfl orens Link & Otto), Begoniaceae, Cestrum begonia # aurantiacum Lindl., Solanaceae, yellow or orange cestrum Bidens elegans (Brongn.) Schltdl. (= C. purpureum (Lindl.) Standl.), Solan- bipinnata L., Asteraceae, Spanish black jack aceae, crimson cestrum biternata (Lour.) Merr. & Sherff, Asteraceae, fi ve-leaved black jack laevigatum Schltdl., Solanaceae, inkberry pilosa L., Asteraceae, black jack parqui L’Hér., Solanaceae, Chilean cestrum sp., Solanaceae Billardiera heterophylla (Lindl.) L.W.Cayzer & Crisp (= Sollya heterophylla Lindl.), Pittosporaceae, bluebell creeper # Chamaesyce prostrata (Aiton) Small (= Euphorbia prostrata Aiton), Euphorbiaceae, Boerhavia erecta L., Nyctaginaceae, erect boerhavia hairy creeping milkweed Bougainvillea glabra Choisy, Nyctaginaceae, bougainvillea, *2004 # serpens (Kunth) Small (= Euphorbia serpens Kunth), Euphorbiaceae, Brachychiton populneus (Schott & Endl.) R.Br., Sterculiaceae, milkweed kurrajong, *2006 # Briza maxima L. (= B. major K.Presl), Poaceae, quaking grass Chenopodium album L., Chenopodiaceae, white goosefoot Chondrilla juncea L., Asteraceae, skeletonweed, *2003 # Bromus Chorizema cordatum Lindl., Fabaceae, Australian fl ame pea # catharticus Vahl (= B. unioloides Kunth, B. willdenowii Kunth), Chromolaena odorata (L.) R.M.King & H.Rob. (= Eupatorium odoratum Poaceae, rescue grass L.), Asteraceae, triffi d weed diandrus Roth, Poaceae, ripgut brome Cichorium intybus L., Asteraceae, chicory pectinatus Thunb. (= B. adoensis Hochst. ex Steud.), Poaceae, Cinnamomum camphora (L.) J.Presl, Lauraceae, camphor tree Japanese brome Cirsium Brugmansia ×candida Pers., (= Datura candida (Pers.) Saff.), Solan- arvense (L.) Scop., Asteraceae, Canada thistle aceae, moonfl ower bush vulgare (Savi) Ten. (= C. lanceolatum (L.) Scop.), Asteraceae, spear Bryophyllum thistle delagoense (Eckl. & Zeyh.) Schinz (= Kalanchoe tubifl ora (Harv.) Cissus antarctica Vent., Vitaceae, kangaroo vine, *2002 # Raym.-Hamet), Crassulaceae, chandelier plant pinnatum (Lam.) Oken (= Kalanchoe pinnata (Lam.) Pers.), Crassu- Citrus laceae, green mother of millions, *2005 ?naturalized limon (L.) Burm.f. (= C. limonum Risso), Rutaceae, lemon proliferum Bowie ex Hook. (= Kalanchoe prolifera (Bowie ex Hook.) sp., Rutaceae Raym.-Hamet, Crassulaceae, *2005 ?naturalized Clusia rosea Jacq., Clusiaceae, balsam fi g/apple, *2003 # Buddleja Coffea arabica L., Rubiaceae, arabica coffee, †Zimbabwe # abundant davidii Franch., Buddlejaceae, Chinese sagewood, *2004 ?naturalized locally ?Buddleja madagascariensis Lam., Buddlejaceae, Madagascar sage- Coix lacryma-jobi L., Poaceae, Job’s tears wood # Colocasia esculenta (L.) Schott, Araceae, elephant’s ear Bothalia 37,2 (2007) 241

APPENDIX 5.—Species checklist (cont.)

Commelina benghalensis L., Commelinaceae, Benghal wandering Jew Eucalyptus Convolvulus arvensis L., Convolvulaceae, fi eld bindweed camaldulensis Dehnh., Myrtaceae, red river gum cinerea F.Muell. ex Benth., Myrtaceae, fl orist’s gum Conyza cladocalyx F.Muell., Myrtaceae, sugar gum bonariensis (L.) Cronquist (= Erigeron bonariensis L.), Asteraceae, cloeziana F.Muell., Myrtaceae, iron gum fl ax-leaf fl eabane conferruminata D.J.Carr & S.G.M.Carr (E. lehmannii (Schauer) canadensis (L.) Cronquist (= Erigeron canadensis L.), Asteraceae, Benth. misapplied in South Africa), Myrtaceae, bald island horseweed fl eabane marlock or ‘spider gum’ primulifolia (Lam.) Cuatrec. & Lourteig (= C. chilensis Spreng.), diversicolor F.Muell., Myrtaceae, karri Asteraceae, Chilean fl eabane ?exserta F.Muell., Myrtaceae, Queensland peppermint sp., Asteraceae fastigata H.Deane & Maiden, Myrtaceae, cut-tail gum sumatrensis (Retz.) E.Walker (= C. albida Willd. ex Spreng.), globulus Labill., Myrtaceae, blue gum Asteraceae, tall fl eabane gomphocephala DC., Myrtaceae, tuart Coreopsis lanceolata L., Asteraceae, tickseed grandis W.Hill ex Maiden (E. saligna Sm. misapplied in South Africa), Myrtaceae, saligna gum Cortaderia leucoxylon F.Muell., Myrtaceae, white ironbark ?# jubata (Lemoine ex Carrière) Stapf, Poaceae, purple Pampas grass microcorys F.Muell., Myrtaceae, tallow gum selloana (Schult.) Asch. & Graebn., Poaceae, common Pampas grass microtheca F.Muell., Myrtaceae, coolabah Corymbia fi cifolia (F.Muell.) K.D.Hill & L.A.S.Johnson (= Eucalyptus paniculata Sm., Myrtaceae, grey ironbark fi cifolia F.Muell.), Myrtaceae, red fl owering gum # regnans F.Muell., Myrtaceae, mountain ash robusta Sm., Myrtaceae, swamp mahogany gum ?# Cosmos bipinnatus Cav. (= Bidens formosa (Bonato) Sch. Bip.), sideroxylon A.Cunn ex Woolls, Myrtaceae, black ironbark ?# Asteraceae, cosmos sp., Myrtaceae tereticornis Sm., Myrtaceae, forest red gum ?# Cotoneaster coriaceus Franch. (= C. lacteus W.W.Sm.), Rosaceae # Eugenia unifl ora L., Myrtaceae, pitanga franchetii Bois, Rosaceae, orange cotoneaster Euphorbia glaucophyllus Franch., Rosaceae, late cotoneaster heterophylla L. (= E. geniculata Ortega), Euphorbiaceae, annual pannosus Franch., Rosaceae, silver-leaf cotoneaster poinsettia sp., Rosaceae leucocephala Lotsy, Euphorbiaceae, white poinsettia, *2005 Crataegus ?naturalized sp., Rosaceae # peplus L., Euphorbiaceae, stinging milkweed ×lavallei Hérincq (= C. carrierei Vauvel ex Carriére), Rosaceae, pulcherrima Willd. ex Klotzsch (= Poinsettia pulcherrima (Willd. ex Lavallee thorn Klotzsch) Graham), Euphorbiaceae, poinsettia ?# Crotalaria agatifl ora Schweinf. subsp. agatifl ora, Fabaceae, canary-bird Euryops chrysanthemoides (DC.) B.Nord., Asteraceae, †Zimbabwe # bush but indigenous in South Africa Cryptomeria japonica (L.f.) D.Don, Cupressaceae, Japanese cedar # Fallopia Cryptostegia grandifl ora R.Br., Asclepiadaceae, rubber vine convolvulus (L.) Á.Löve (= Bilderdykia convolvulus (L.) Dumort), Cuphea ignea A.DC., Lythraceae, cigarette bush # Polygonaceae, climbing knotweed Cupressus sachalinensis (F.Schmidt) Ronse Decr. (= Polygonum sachalinense arizonica Greene (= C. glabra Sudw.), Cupressaceae, Arizona cypress F.Schmidt, Reynoutria sachalinensis (F.Schmidt) Nakai), Poly- lusitanica Mill. (= C. lindleyi Klotzsch ex Endl.), Cupressaceae, Mexi- gonaceae, giant knotweed, *2005 (PRE 1980) naturalized can cypress Ficus sp., Cupressaceae carica L., Moraceae, fi g elastica Roxb. ex Hornem.(= F. decora hort.), Moraceae, rubber fi g # Cuscuta macrophylla Desf. ex Pers., Moraceae, Australian banyan # campestris Yunck., Convolvulaceae, common dodder pumila L., Moraceae, tickey creeper suaveolens Ser., Convolvulaceae, lucerne dodder Flaveria bidentis (L.) Kuntze (= F. contrayerba (Cav.) Pers.), Asteraceae, Cyathea cooperi (Hook. ex F.Muell.) Domin (= Sphaeropteris cooperi smelter’s bush (Hook. ex F. Muell.) R.M.Tryon), Cyatheaceae, Australian tree Foeniculum vulgare Mill., Apiaceae, fennel fern, *2005 # Fraxinus Cydonia oblonga Mill.( = C. vulgaris Pers.), Rosaceae, quince americana L., Oleaceae, American ash Cytisus scoparius (L.) Link (= Genista scoparia (L.) Lam.), Fabaceae, angustifolia Vahl, Oleaceae, Algerian ash Scotch broom sp., Oleaceae Dahlia spp., Asteraceae, garden dahlias ?naturalized Fuchsia sp., Onagraceae, fuchsia # Datura Fumaria muralis Sond. ex Koch, Fumariaceae, wall fumitory, *2001 # ferox L., Solanaceae, large thorn apple Genista monspessulana (L.) L.A.S.Johnson (= Cytisus candicans (L.) innoxia Mill. (D. metel L. misapplied in South Africa), Solanaceae, DC., C. monspessulanus L.), Fabaceae, Montpellier broom downy thorn apple sp., Solanaceae Glandularia stramonium L., Solanaceae, common thorn apple aristigera (S.Moore) Tronc.(= Verbena tenuisecta Briq.), Verbenaceae, fi ne-leaved verbena Delonix regia (Bojer ex Hook.) Raf. (= Poinciana regia Bojer ex Hook.), ×hybrida (hort. ex Groenl. & Rümpler) G.L.Nesom & Pruski (= Verbena Fabaceae, fl amboyant ×hybrida hort. ex Groenl. ex Rümpler), Verbenaceae, garden Desmanthus virgatus (L.) Willd. (= D. depressus Humb. & Bonpl. ex verbena ?# Willd.), Fabaceae, ground tamarind Glebionis coronaria (L.) Cass. ex Spach (= Chrysanthemum coronarium Desmodium uncinatum (Jacq.) DC., Fabaceae, silverleaf desmodium, L.), Asteraceae, chrysanthemum greens †Zimbabwe # abundant locally Gleditsia triacanthos L., Fabaceae, honey locust Dracocephalum canariense L. (= Cedronella canariensis (L.) Webb & Glyceria maxima (Hartm.) Holmb. (= G. aquatica (L.) Wahlb., Poa Berthel.), Lamiaceae, hortela de burro aquatica L.), Poaceae, reed meadow grass, *2002 # Duranta erecta L. (= D. repens L., D. plumieri Jacq.), Verbenaceae, Gmelina arborea Roxb., Verbenaceae, white teak, †Malawi # abundant forget-me-not-tree locally Dysphania ambrosioides (L.) Mosyakin & Clemants (= Chenopodium Gnaphalium luteoalbum L. (= Pseudognaphalium luteoalbum (L.) ambrosioides L.), Chenopodiaceae, American goosefoot Hilliard & B.L.Burtt), Asteraceae, Jersey cudweed Echinopsis spachiana (Lem.) Friedrich & G.D.Rowley (= Trichocereus Gomphrena celosioides Mart. (= G. decumbens Jacq.), Amaranthaceae, spachianus (Lem.) Riccob.), Cactaceae, torch cactus prostrate globe amaranth Echium Grevillea plantagineum L. (= E. lycopsis L.), Boraginaceae, Patterson’s curse banksii R.Br., Proteaceae, Bank’s grevillea, *2004 # very abundant vulgare L., Boraginaceae, blue echium locally Egeria densa Planch. (= Elodea densa (Planch.) Casp.), Hydrocharitaceae, robusta A.Cunn. ex R.Br., Proteaceae, Australian silky oak dense water weed rosmarinifolia A.Cunn., Proteaceae # Eichhornia crassipes (Mart.) Solms, Pontederiaceae, water hyacinth sericea (Sm.) R.Br., Proteaceae, pink spider fl ower # Eragrostis pilosa (L.) P.Beauv., Poaceae, Indian love grass Guilleminea densa (Humb. & Bonpl. ex Schult.) Moq. (= Brayulinea Eriobotrya japonica (Thunb.) Lindl., Rosaceae, loquat densa (Willd.) Small), Amaranthaceae, carrot weed 242 Bothalia 37,2 (2007)

APPENDIX 5.—Species checklist (cont.)

Hakea sinense Lour., Oleaceae, Chinese privet drupacea (C.F.Gaertn.) Roem. & Schult. (= H. suaveolens R.Br.), sp., Oleaceae Proteaceae, sweet hakea vulgare L., Oleaceae, common privet gibbosa (Sm.) Cav., Proteaceae, rock hakea salicifolia (Vent.) B.L.Burtt (= H. saligna (Andrews) Knight), Proteaceae, Lilium formosanum Wallace (= L. longifl orum Thunb. var. formosanum willow hakea Baker, L. phillipinense Baker), Liliaceae, Saint Joseph’s lily sericea Schrad. & J.C.Wendl., Proteaceae, silky hakea Limonium sinuatum (L.) Mill. (= Statice sinuata L.), Plumbaginaceae, victoriae J.Drumm., Proteaceae # statice Harrisia martinii (Labour.) Britton & Rose (= Eriocereus martinii Linaria (Labour.) Riccob.), Cactaceae, harrisia genistifolia (L.) Mill. (= L. dalmatica (L.) Mill.), Scrophulariaceae, Hedera helix L. subsp. canariensis (Willd.) Cout., Araliaceae, Algerian yellow linaria or Canary ivy, *2003 # maroccana Hook.f., Scrophulariaceae, baby snapdragon Hedychium Litsea glutinosa (Lour.) C.B.Rob. (= L. sebifera Pers.), Lauraceae, coccineum Buch.-Ham. ex Sm., Zingiberaceae, red ginger lily Indian laurel coronarium J.König, Zingiberaceae, white ginger lily Lonicera japonica Thunb. ‘Halliana’, Caprifoliaceae, Japanese honey- fl avescens Carey ex Roscoe, Zingiberaceae, yellow ginger lily suckle gardnerianum Sheppard ex Ker Gawl., Zingiberaceae, kahili ginger Lygodium japonicum (Thunb.) Sw., Schizaeaceae, Japanese climbing lily fern # sp., Zingiberaceae Lythrum salicaria L., Lythraceae, purple loosestrife Macfadyena unguis-cati (L.) A.H.Gentry, Bignoniaceae, cat’s claw Helianthus annuus L., Asteraceae, common sunfl ower creeper Heliotropium amplexicaule Vahl, Boraginaceae, blue heliotrope Maireana brevifolia (R.Br.) Paul G.Wilson (= Kochia brevifolia R.Br.), Hibiscus trionum L., Malvaceae, bladderweed Chenopodiaceae, small-leaf bluebush # Homalanthus populifolius Graham, Euphorbiaceae, Queensland poplar Hordeum murinum L., Poaceae, wild barley Malus pumila Mill. var. paradisiaca C.K.Schneid., Rosaceae, paradise Hydrilla verticillata (L.f.) Royle, Hydrocharitaceae, hydrilla, *2006 apple ?# (PRE 1963 but misidentifi ed) naturalized and very abundant at Malva Pongolapoort Dam, KwaZulu-Natal dendromorpha M.F.Ray (= Lavatera arborea L.), Malvaceae, tree Hydrocotyle ranunculoides L.f., Apiaceae, †Zimbabwe # abundant mallow locally linnaei M.F.Ray (= Lavatera cretica L.), Malvaceae, Cretan holly- Hylocereus undatus (Haw.) Britton & Rose, Cactaceae, night-blooming hock cereus parvifl ora L., Malvaceae, small mallow Hypericum Malvastrum coromandelianum (L.) Garcke, Malvaceae, prickly patulum Thunb. (= H. patulum var. forrestii Chitt.), Clusiaceae malvastrum perforatum L., Clusiaceae, St. John’s wort Mangifera indica L., Anacardiaceae, mango Hypochaeris radicata L., Asteraceae, hairy wild lettuce Manihot Hypoestes phyllostachya Baker, Acanthaceae, polka-dot-plant, *2002 # esculenta Crantz (= M. utilissima Pohl), Euphorbiaceae, bitter Ipomoea cassava alba L., Convolvulaceae, moonfl ower grahamii Hook. (= M. dulcis (J.F.Gmel.) Pax var. multifi da (Graham) carnea Jacq. subsp. fi stulosa (Mart. ex Choisy) D.F.Austin (= I. Pax), Euphorbiaceae, hardy cassava # fi stulosa Mart. ex Choisy), Convolvulaceae, potato bush Medicago sativa L. (= M. falcata L.), Fabaceae, alfalfa indica (Burm.) Merr. (= I. congesta R.Br.), Convolvulaceae, perennial morning glory Melaleuca nil (L.) Roth, Convolvulaceae, Japanese morning glory hypericifolia Sm., Myrtaceae, red-fl owering tea tree purpurea (L.) Roth, Convolvulaceae, common morning glory wilsonii F.Muell., Myrtaceae, violet honey-myrtle # sp., Convolvulaceae Melia azedarach L., Meliaceae, seringa or ‘syringa’ Iris pseudacorus L., Iridaceae, yellow fl ag, *2004 # Melilotus alba Medik., Fabaceae, white sweet clover Jacaranda mimosifolia D.Don, Bignoniaceae, jacaranda Metasequoia glyptostroboides Hu & W.C.Cheng, Cupressaceae, dawn Jasminum redwood # humile L., Oleaceae, yellow bush jasmine Metrosideros excelsa Sol. ex Gaertn. (= M. tomentosa A. Rich.), mesnyi Hance, Oleaceae, primrose jasmine ?# Myrtaceae, New Zealand bottlebrush polyanthum Franch., Oleaceae, creeping jasmine, *2001 # Michelia champaca L., Magnoliaceae, champac magnolia, †Zimbabwe # Jatropha Mimosa curcas L., Euphorbiaceae, physic nut, *2005 (1979 in Wells et al. pigra L., Fabaceae, giant sensitive plant (1986)) pudica L. var. hispida Brenan, Fabaceae, sensitive plant gossypiifolia L., Euphorbiaceae, coral plant Mirabilis jalapa L., Nyctaginaceae, four-o’clock sp., Euphorbiaceae Momordica charantia L., Cucurbitaceae, bitter cucumber Juniperus Mondia whitei (Hook.f.) Skeels (= Chlorocodon whitei Hook.f.), Apocyn- pinchotii Sudw., Cupressaceae, red-berry juniper # aceae, †Zimbabwe & Malawi # but indigenous in South Africa sp., Cupressaceae Monstera deliciosa Liebm., Araceae, Swiss-cheese plant # virginiana L., Cupressaceae, red cedar Montanoa hibiscifolia Benth., Asteraceae, tree daisy Moringa oleifera Lam. (= M. pterygosperma Gaertn.), Moringaceae, Koelreuteria paniculata Laxm. (= K. apiculata Rehder & E.H.Wilson), horse-radish tree # Sapindaceae, golden-rain tree *2001 # Morus alba L., Moraceae, white or common mulberry Lactuca serriola L. (= L. scariola L.), Asteraceae, wild lettuce Musa sp., Musaceae, banana tree # Lagerstroemia indica L., Lythraceae, pride-of-India Murraya paniculata (L.) Jack. (= M. exotica L. ), Rutaceae, orange Lantana camara L., Verbenaceae, lantana jessamine, *2005 # Lemna Myoporum tenuifolium G.Forst. subsp. montanum (R.Br.) Chinnock (= gibba L., Lemnaceae, duckweed M. montanum R.Br.)(M. acuminatum R.Br. misapplied in South sp., Lemnaceae Africa), Myoporaceae, manatoka Lepidium Myriophyllum didymum L. (= Coronopus didymus (L.) Sm.), Brassicaceae, aquaticum (Vell.) Verdc. (= M. brasiliense Cambess.), Haloragaceae, swinecress parrot’s feather draba L. (= Cardaria draba (L.) Desv.), Brassicaceae, hoary cardaria spicatum L., Haloragaceae, spiked water-milfoil Leptospermum laevigatum (Gaertn.) F.Muell., Myrtaceae, Australian Nassella myrtle tenuissima (Trin.) Barkworth (= Stipa tenuissima Trin.), Poaceae, white Leucaena leucocephala (Lam.) de Wit (= L. glauca Benth.), Fabaceae, tussock leucaena trichotoma (Nees) Hack. ex Arechav. (= Stipa trichotoma Nees), Poaceae, nassella tussock Ligustrum japonicum Thunb., Oleaceae, Japanese wax-leaved privet Nasturtium offi cinale R.Br. (= Rorippa nasturtium-aquaticum (L.) lucidum W.T.Aiton, Oleaceae, Chinese wax-leaved privet Hayek), Brassicaceae, watercress ovalifolium Hassk., Oleaceae, Californian privet Nephrolepis exaltata (L.) Schott, Nephrolepidaceae, sword fern Bothalia 37,2 (2007) 243

APPENDIX 5.—Species checklist (cont.)

Nerium oleander L., Apocynaceae, oleander Physalis Nicandra physalodes (L.) Gaertn., Solanaceae, apple-of-Peru peruviana L., Solanaceae, Cape gooseberry viscosa L., Solanaceae, sticky gooseberry Nicotiana glauca Graham, Solanaceae, wild tobacco Phytolacca tabacum L., Solanaceae, tobacco ?# dioica L., Phytolaccaceae, belhambra icosandra L. (= P. octandra L.), Phytolaccaceae, forest inkberry Nymphaea mexicana Zucc., Nymphaeaceae, yellow waterlily Pinus ×marliacea W.Watson, Nymphaeaceae, Marliac hybrid waterlily canariensis C.Sm., Pinaceae, Canary pine elliottii Engelm., Pinaceae, slash pine Oenothera halepensis Mill., Pinaceae, Aleppo pine biennis L., Onagraceae, evening primrose patula Schiede ex Schltdl. & Cham., Pinaceae, patula pine glazioviana Micheli (= O. erythrosepala Borbás), Onagraceae, evening pinaster Aiton, Pinaceae, cluster pine primrose pinea L., Pinaceae, umbrella pine indecora Cambess., Onagraceae, evening primrose radiata D.Don, Pinaceae, radiata pine jamesii Torr. & A.Gray, Onagraceae, giant evening primrose roxburghii Sarg. (= P. longifolia Roxb. ex Lamb.), Pinaceae, chir pine laciniata Hill, Onagraceae, cutleaf evening primrose sp., Pinaceae rosea L’Hér. ex Aiton, Onagraceae, rose evening primrose taeda L., Pinaceae, loblolly pine sp., Onagraceae tetraptera Cav., Onagraceae, white evening primrose Pistia stratiotes L., Araceae, water lettuce Pittosporum undulatum Vent., Pittosporaceae, Australian cheesewood Olyra latifolia L., Poaceae Pityrogramma calomelanos (L.) Link, Adiantaceae, golden fern Opuntia Plantago aurantiaca Lindl., Cactaceae, jointed cactus lanceolata L., Plantaginaceae, narrow-leaved ribwort engelmannii Salm-Dyck ex Engelm. (= O. lindheimeri Engelm.), major L., Plantaginaceae, broad-leaved ribwort Cactaceae, small round-leaved prickly pear virginica L., Plantaginaceae, dwarf plantain exaltata A.Berger (= Austrocylindropuntia exaltata (A.Berger) Backeb.), Cactaceae, long-spine cactus Platanus fi cus-indica (L.) Mill. (= O. megacantha Salm-Dyck), Cactaceae, sp., Platanaceae # sweet prickly pear ×acerifolia (Aiton) Willd. (= P. hispanica auct.), Platanaceae, London fulgida Engelm. (= Cylindropuntia fulgida (Engelm.) F.M.Knuth)(O. planetree, *2004 # rosea DC. and Cylindropuntia rosea (DC.) Backeb. misapplied in South Africa), Cactaceae, chainfruit cholla or ‘rosea cactus’ Plectranthus comosus Sims (= Coleus grandis Cramer)(P. barbatus humifusa (Raf.) Raf. (= O. compressa auct.), Cactaceae, large-fl owered Andrews misapplied in South Africa), Lamiaceae, Abyssinian coleus prickly pear Polygonum aviculare L., Polygonaceae, prostrate knotweed imbricata (Haw.) DC. (= Cylindropuntia imbricata (Haw.) F.M.Knuth), Polypogon monspeliensis (L.) Desf., Poaceae, beardgrass Cactaceae, imbricate prickly pear Pomaderris kumeraho A.Cunn., Rhamnaceae, kumarahou # microdasys (Lehm.) Pfeiff., Cactaceae, yellow bunny-ears Pontederia cordata L., Pontederiaceae, pickerel weed monacantha Haw. (= O. vulgaris auct.), Cactaceae, cochineal prickly Populus pear alba L., Salicaceae, white poplar robusta H.L.Wendl. ex Pfeiff., Cactaceae, blue-leaf cactus deltoides W.Bartram ex Marshall, Salicaceae, match poplar sp., Cactaceae nigra L. var. italica Münchh., Salicaceae, Lombardy poplar spinulifera Salm-Dyck, Cactaceae, large round-leaved prickly pear ?# ×canescens (Aiton) Sm., Salicaceae, grey poplar stricta (Haw.) Haw. (possibly both var. dillenii and var. stricta), Cac- taceae, Australian pest pear Portulaca oleracea L., Portulacaceae, purslane stricta ×humifusa?, Cactaceae Prosopis tomentosa Salm-Dyck, Cactaceae, velvet opuntia, *2003 # glandulosa Torr. var. torreyana (Benson) Johnst., Fabaceae, honey Orobanche minor Sm., Orobanchaceae, clover broomrape mesquite Oxalis corniculata L., Oxalidaceae, creeping oxalis velutina Wooton, Fabaceae, velvet mesquite Pandanus sp., Pandanaceae, screw-pine # Prunus Paraserianthes lophantha (Willd.) I.C.Nielsen (= Albizia lophantha armeniaca L., Rosaceae, apricot (Willd.) Benth.), Fabaceae, stinkbean cerasoides D.Don, Rosaceae, Himalayan fl owering cherry, †Zimbabwe Parkinsonia aculeata L., Fabaceae, Jerusalem thorn # abundant locally Parthenium hysterophorus L., Asteraceae, parthenium persica (L.) Batsch, Rosaceae, peach Parthenocissus quinquefolia (L.) Planch., Vitaceae, Virginia cree- serotina Ehrh., Rosaceae, black cherry per # Psidium Paspalum cattleianum Sabine (= P. littorale Raddi var. longipes (O.Berg) dilatatum Poir., Poaceae, common paspalum Fosberg), Myrtaceae, strawberry guava notatum Flüggé, Poaceae, *2006 (PRE 1944) guajava L., Myrtaceae, guava quadrifarium Lam., Poaceae guineense Sw., Myrtaceae, Brazilian guava urvillei Steud., Poaceae, tall paspalum sp., Myrtaceae Passifl ora ×durbanensis Baijnath ined., Myrtaceae, Durban guava caerulea L., Passifl oraceae, blue passion fl ower Pterocarya stenoptera C.DC., Juglandaceae, Chinese wing-nut # edulis Sims, Passifl oraceae, purple granadilla Pueraria montana (Lour.) Merr. var. lobata (Willd.) Maesen & sp., Passifl oraceae S.M.Almeida (= P. lobata (Willd.) Ohwi), Fabaceae, kudzu vine suberosa L., Passifl oraceae, devil’s pumpkin Punica granatum L., Punicaceae, pomegranate subpeltata Ortega, Passifl oraceae, granadina tripartita (Juss.) Poir. var. mollissima (Kunth) Holm-Niels. & P.Jorg. Pyracantha (= P. mollissima (Kunth) L.H.Bailey), Passifl oraceae, banana poka angustifolia (Franch.) C.K.Schneid., Rosaceae, yellow fi rethorn coccinea M.Roem., Rosaceae, red fi rethorn Pennisetum crenulata (D.Don) M.Roem., Rosaceae, Himalayan fi rethorn clandestinum Hochst. ex Chiov., Poaceae, Kikuyu grass sp., Rosaceae purpureum Schumach., Poaceae, Napier grass setaceum (Forssk.) Chiov., Poaceae, fountain grass Pyrus sp., Rosaceae, pear tree ?# sp., Poaceae Quercus villosum R.Br. ex Fresen., Poaceae, feathertop canariensis Willd. (= Q. mirbeckii Durieu), Fagaceae, Algerian oak ?# Pereskia aculeata Mill., Cactaceae, pereskia cerris L., Fagaceae, Turkey oak ?# Persea americana Mill. (= P. gratissima C.F.Gaertn.), Lauraceae, palustris Münchh., Fagaceae, pin oak avocado pear # robur L., Fagaceae, English oak Persicaria lapathifolia (L.) Gray (= Polygonum lapathifolium L.), sp., Fagaceae Polygonaceae, spotted knotweed suber L., Fagaceae, cork oak ?# Phoenix Quisqualis indica L., Combretaceae, Rangoon creeper *2006 # canariensis Hort. ex Chabaud, Arecaceae, Canary date palm dactylifera L., Arecaceae, real date palm Richardia brasiliensis Gomes, Rubiaceae, tropical richardia Phormium tenax J.R.Forst. & G.Forst., Phormiaceae, New Zealand fl ax # humistrata Steud., Rubiaceae, Peelton richardia 244 Bothalia 37,2 (2007)

APPENDIX 5.—Species checklist (cont.)

Ricinus communis L., Euphorbiaceae, castor-oil plant tree tomato Rivina humilis L., Phytolaccaceae, bloodberry capsicoides All., Solanaceae, devil’s apple Robinia pseudoacacia L., Fabaceae, black locust chrysotrichum Schltdl. (= S. hispidum auctt. non Pers.), Solanaceae, giant devil’s fi g Rosa elaeagnifolium Cav., Solanaceae, silver-leaf bitter apple multifl ora Thunb., Rosaceae, multifl ora rose mauritianum Scop. (= S. auriculatum Aiton), Solanaceae, bugweed rubiginosa L. (= R. eglanteria L.), Rosaceae, eglantine pseudocapsicum L., Solanaceae, Jerusalem cherry sp., Rosaceae # seaforthianum Andrews, Solanaceae, potato creeper ?×odorata (Andrews) Sweet, Rosaceae, tea rose # sisymbriifolium Lam., Solanaceae, dense-thorned bitter apple Rubus sp., Solanaceae cuneifolius Pursh, Rosaceae, American bramble torvum Sw. (= S. mannii C.H.Wright), Solanaceae fl agellaris Willd., Rosaceae viarum Dunal, Solanaceae, tropical soda apple, *2006 (PRE 1962) fruticosus L. agg., Rosaceae, European blackberry Sonchus oleraceus L., Asteraceae, sowthistle ?pascuus L.H.Bailey, Rosaceae Sophora cf. davidii (Franch.) Skeels, Fabaceae # phoenicolasius Maxim., Rosaceae Sorghum halepense (L.) Pers. (= S. almum Parodi), Poaceae, Johnson rosifolius Sm., Rosaceae grass sp., Rosaceae Spartium junceum L., Fabaceae, Spanish broom ×proteus C.H.Stirt., Rosaceae, Bramble, Graskop/Sabie hybrid Spathodea campanulata P.Beauv., Bignoniaceae, African fl ame tree Rumex Sphagneticola trilobata (L.) Pruski (= Thelechitonia trilobata (L.) acetosella L. subsp. pyrenaicus (Pourr. ex Lapeyr.) Akeroyd (= R. H.Rob. & Cuatrec, Wedelia trilobata (L.) Hitchc.), Asteraceae, angiocarpus auct.), Polygonaceae, sheep sorrel Singapore daisy crispus L., Polygonaceae, curly dock Spiraea cantoniensis Lour., Rosaceae, Cape may # usambarensis (Dammer) Dammer (= R. nervosus Vahl var. usambarensis Stellaria media (L.) Vill., Caryophyllaceae, chickweed Dammer), Polygonaceae, rumex Stenocarpus sinuatus Endl., Proteaceae, fi rewheel tree ?# Styphnolobium japonicum (L.) Schott (= Sophora japonica L.), Saccharum offi cinarum L., Poaceae, sugar cane Fabaceae, Japanese pagoda tree ?# Salix Symphyotrichum subulatum (Michx.) G.L.Nesom var. squamatum babylonica L., Salicaceae, weeping willow (Spreng.) S.D.Sundb. (= Aster squamatus (Spreng.) Hieron.), caprea L., Salicaceae, pussy willow Asteraceae, swamp aster fragilis L., Salicaceae, crack willow Syncarpia glomulifera (Sm.) Nied. (= S. laurifolia Ten.), Myrtaceae, turpentine tree Salsola kali L., Chenopodiaceae, common saltwort Syzygium tragus L. (= S. australis R.Br.), Chenopodiaceae, Russian tumbleweed cumini (L.) Skeels, Myrtaceae, jambolan jambos (L.) Alston, Myrtaceae, rose apple Salvia tiliifolia Vahl, Lamiaceae, Lindenleaf sage, *2005 (PRE 1943) paniculatum Gaertn. (= Eugenia myrtifolia Sims), Myrtaceae, Salvinia molesta D.S.Mitch. (S. auriculata Aubl. misapplied in South Australian water pear Africa), Salviniaceae, Kariba weed Tabebuia chrysotricha (Mart. ex DC.) Standl. (= Tecoma chrysotricha Sambucus Mart. ex DC.), Bignoniaceae, yellow trumpet tree ?# canadensis L. (= S. nigra L. subsp. canadensis (L.) Bolli), Capri- Tagetes minuta L., Asteraceae, khaki weed foliaceae, Canadian elder nigra L., Caprifoliaceae, European elder, *2004 Tamarix sp., Caprifoliaceae chinensis Lour., Tamaricaceae, Chinese tamarisk ramosissima Ledeb., Tamaricaceae, pink tamarisk Scheffl era sp., Tamaricaceae actinophylla (Endl.) Harms (= Brassaia actinophylla Endl.), Araliaceae, Australian cabbage tree # Taraxacum offi cinale F.H.Wigg. agg., Asteraceae, common dandelion arboricola (Hayata) Merr., Araliaceae, dwarf umbrella tree, *2005 # Tecoma elegantissima (hort. Veitch. ex Mast.) Lowry & Frodin (= Dizygotheca stans (L.) Juss. ex Kunth, Bignoniaceae, yellow bells elegantissima (hort. Veitch. ex Mast.) R.Vig. Guillaumin), tenuifl ora (A.DC.) Fabris, Bignoniaceae, *2004 Araliaceae, *2005 # Tephrocactus Schinus articulatus (Pfeiff.) Backeb., Cactaceae, paper-spine cholla molle L., Anacardiaceae, pepper tree ?aoracanthus (Lem.) Lem. (= ?Opuntia aoracantha Lemaire), Cactaceae terebinthifolius Raddi, Anacardiaceae, Brazilian pepper tree sp., Cactaceae Schizolobium parahyba (Vell.) S. F.Blake var. parahyba (= S. excelsum Thevetia peruviana (Pers.) K.Schum. (= T. neriifolia Juss. ex Steud.), Vogel), Fabaceae, parasol tree # Apocynaceae, yellow oleander Schkuhria pinnata (Lam.) Kuntze ex Thell., Asteraceae, dwarf marigold Tipuana tipu (Benth.) Kuntze (= T. speciosa Benth.), Fabaceae, tipu tree Senna bicapsularis (L.) Roxb. (= Cassia bicapsularis L.), Fabaceae, rambling Tithonia cassia diversifolia (Hemsl.) A.Gray, Asteraceae, Mexican sunfl ower corymbosa (Lam.) H.S.Irwin & Barneby (= Cassia corymbosa Lam.), rotundifolia (Mill.) S.F.Blake, Asteraceae, red sunfl ower Fabaceae, autumn cassia Toona ciliata M.Roem. (= Cedrela toona Roxb. ex Willd.), Meliaceae, didymobotrya (Fresen.) H.S.Irwin & Barneby (= Cassia didymobotrya toon tree Fresen.), Fabaceae, peanut butter cassia Torilis arvensis (Huds.) Link., Apiaceae, spreading hedge-parsley hirsuta (L.) H.S.Irwin & Barneby (= Cassia hirsuta L.), Fabaceae Toxicodendron succedaneum (L.) Kuntze (= Rhus succedanea L.), Ana- multiglandulosa (Jacq.) H.S.Irwin & Barneby (= Cassia multiglan- cardiaceae, wax tree dulosa Jacq., C. tomentosa L.f.), Fabaceae obtusifolia (L.) H.S.Irwin & Barneby (= Cassia obtusifolia L.), Fabaceae Tradescantia occidentalis (L.) Link (= Cassia occidentalis L.), Fabaceae, wild coffee zebrina hort. ex Bosse (= Zebrina pendula Schnizl.), Commelinaceae, pendula (Willd.) H.S.Irwin & Barneby var. glabrata (Vogel) H.S.Irwin wandering jew, *2005 # & Barneby (= Cassia coluteoides Collad.), Fabaceae fl uminensis Vell., Commelinaceae, wandering Jew, *2001 (Wells et septemtrionalis (Viv.) H.S.Irwin & Barneby (= Cassia fl oribunda sensu al. 1986) Brenan non Cav.), Fabaceae, arsenic bush Tragopogon dubius Scop. (= T. major Jacq.), Asteraceae, yellow salsify sp., Fabaceae Tridax procumbens L., Asteraceae, tridax daisy Sesbania Triplaris americana L., Polygonaceae, triplaris ?# bispinosa (Jacq.) W.Wight var. bispinosa (= S. aculeata Pers.), Fabaceae, Triticum aestivum L., Poaceae, volunteer wheat spiny sesbania Tropaeolum majus L., Tropaeolaceae, nasturtium punicea (Cav.) Benth., Fabaceae, red sesbania Ulex europaeus L., Fabaceae, European gorse Sigesbeckia orientalis L., Asteraceae, St. Paul’s wort Ulmus Silybum marianum (L.) Gaertn., Asteraceae, milk thistle parvifolia Jacq. (= U. chinensis Pers.), Ulmaceae, Chinese elm Sisymbrium orientale L., Brassicaceae, Indian hedge mustard procera Salisb., Ulmaceae, English elm # Sisyrinchium sp., Iridaceae sp., Ulmaceae # Solanum Verbena betaceum Cav. (= Cyphomandra betacea (Cav.) Sendtn.), Solanaceae, bonariensis L., Verbenaceae, wild verbena Bothalia 37,2 (2007) 245

APPENDIX 5.—Species checklist (cont.)

Verbena (cont.) trifolia L., Verbenaceae, Indian three-leaf vitex, *2004 # brasiliensis Vell., Verbenaceae, slender wild verbena sp., Vitaceae, grape # offi cinalis L., Verbenaceae, European verbena rigida Spreng. var. rigida (= V. venosa Gillies & Hook.), Verbenaceae, Washingtonia sp., Arecaceae, petticoat palm ?# veined vervain Wigandia urens (Ruiz & Pav.) Kunth var. caracasana (Kunth) D.N.Gibson Verbesina encelioides (Cav.) Benth. & Hook.f. ex A.Gray, Asteraceae, (= W. caracasana Kunth), Hydrophyllaceae, wigandia # golden crownbeard Wisteria fl oribunda (Willd.) DC., Fabaceae, Japanese wisteria # Vinca major L., Apocynaceae, greater periwinkle Viola Xanthium hederacea Labill. (= Erpetion reniforme Sweet), Violaceae, Australian sp., Asteraceae violet, *2001 # spinosum L., Asteraceae, spiny cocklebur priceana Pollard (= V. sororia Willd.), Violaceae, confederate violet, strumarium L., Asteraceae, large cocklebur *2005 # Vitex Yucca aloifolia L., Agavaceae, Spanish bayonet agnus-castus L., Verbenaceae, lilac chastetree, *2004 (PRE 1975) Zinnia peruviana (L.) L. (= Z. multifl ora L.), Asteraceae, redstar zinnia

APPENDIX 6.—Characteristics of prominent invaders in study area

Plant name Family Origin GF W LC P SR VR Disp. agent Cult. use VC Acacia cyclops Fabaceae ST (Aus.) t/s woo per ev seed bir, mam #c/b, bar sa, fy, sk dealbata Fabaceae ST (Aus.) t woo per ev seed cop wat, ?ant, silc, #bar, sa, fo, gr ?bir orn decurrens Fabaceae ST (Aus.) t woo per ev seed cop wat, ?ant, silc, #bar, gr ?bir orn longifolia Fabaceae ST (Aus.) t/s woo per ev seed wat, bir, ant #c/b, bar, orn fy, fo mearnsii Fabaceae ST (Aus.) t woo per ev seed cop wat, ?ant, #silc, bar, sa, fy, fo, gr, sk ?bir orn melanoxylon Fabaceae ST (Aus.) t woo per ev seed suc bir, wat #silc, bar, fy, fo orn pycnantha Fabaceae ST (Aus.) t woo per ev seed wat, mam, #silc, c/b, fy ?bir bar, orn saligna Fabaceae ST (Aus.) t/s woo per ev seed cop wat, mam, silc, agrc, #c/ sa, fy, fo, sk ant b, bar, orn Achyranthes aspera Amaranthaceae T (?Afr.) h her per ev/d seed mam none fo Agave americana var. america- Agavaceae T (Am.) s suc per ev seed suc win, hum orn, #bar, sa, nk na agrc sisalana Agavaceae T (Am.) s suc per ev suc, bul wat, hum bar, #agrc, sa orn Ageratum conyzoides Asteraceae T (Am.) h her ann germ seed win #orn sa, fo houstonianum Asteraceae T (Am.) h her ann germ seed win #orn sa, fo Argemone mexicana Papaveraceae T (Am.) h her ann germ seed wat, ?hum none sa (soil),?ant ochroleuca subsp. ochro- Papaveraceae T (Am.) h her ann germ seed wat, ?hum none sa leuca (soil),?ant Arundo donax Poaceae NT (Med. & g/r sem per ev rhz, div wat, hum #agrc, orn, sa, fy, gr, Asia) bar nk, sk Atriplex infl ata Chenopodiaceae ST (Aus.) h sem per ev/d seed win #?agrc nk, sk nummularia subsp. Chenopodiaceae ST (Aus.) s woo/ per ev/d seed win #agrc, bar nk, sk nummularia sem Azolla fi liculoides Azollaceae T (Am.) h her var var spore div wat, bir #orn gr, nk Caesalpinia decapetala Fabaceae Tl (Asia) s/c woo per ev seed wat, ?hum, #bar, orn sa, fo, gr mam (cattle) Cardiospermum grandifl orum Sapindaceae T (Am.) c sem per ev/d seed wat, win #orn sa, fo halicacabum Sapindaceae T (?Am.) c sem per ev/d seed wat, win #orn sa

Origin: ST, southern temperate, south of or straddling Tropic of Capricorn; NT, northern temperate, north of or straddling Tropic of Cancer; T, tropical, between or straddling Tropics of Capricorn and Cancer. GF, growth form: tree; tree/shrub; shrub; herb; herb/shrub; grass; grass/reed; climber; shrub/climber. W, woodiness: woody; semi-woody; herbaceous; succulent. LC, life cycle: perennial, annual, variable, biennial. P, perennation: evergreen; deciduous; evergreen/deciduous; variable; germinative. SR, sexual reproduction: seed, spore. VR, vegetative reproduction: coppice; sucker; division; rhz, rhizome; bulbil; stolon; tuber; runner. Dispersal agent: wind; water; bird; mammal; human; ant. Cultivated use: # primary (= major) use; ornamental; barrier; cover/binder; agricultural crop; silvicultural crop; none. VC, vegetation category: savanna biome; fynbos biome; forest habitats; grassland biome; nama-karoo biome; succulent karoo biome. Afr., Africa; Am., America; Aus., Australia; Eur., Europe; Med., Mediterranean. 246 Bothalia 37,2 (2007)

APPENDIX 6.—Characteristics of prominent invaders in study area (cont.)

Plant name Family Origin GF W LC P SR VR Disp. agent Cult. use VC Casuarina cunninghamiana Casuarinaceae T (Aus.) t woo per ev seed wat, win orn, c/b, #bar fo equisetifolia Casuarinaceae T (Pantrop.) t woo per ev seed wat, win orn, #c/b, bar fo Cereus jamacaru Cactaceae T (Am.) t/s suc per ev seed div bir #orn, bar sa Cestrum laevigatum Solanaceae T (Am.) t/s woo per ev seed bir #orn, bar sa, fo Chromolaena odorata Asteraceae T (Am.) s woo per ev seed win #orn sa, fo Cinnamomum camphora Lauraceae NT (Asia) t woo per ev seed bir #orn, silc, fo agrc Cirsium vulgare Asteraceae NT (Eur., N h her bie germ seed win none gr Afr. & Asia) Datura ferox Solanaceae T (Am.) h her ann germ seed wat, ?hum none sa, gr (soil), ?ant innoxia Solanaceae T(Am.) h her var var seed wat, ?hum none sa, gr (soil), ?ant stramonium Solanaceae T (Am.) h her ann germ seed wat, ?hum #agrc sa, gr (soil), ?ant Eichhornia crassipes Pontederiaceae T (Am.) h her per ev seed div wat, hum #orn sa, fy, gr Eucalyptus camaldulensis Myrtaceae T (Aus.) t woo per ev seed cop wat, win #silc, bar, fy, fo, sk orn, agrc diversicolor Myrtaceae ST (Aus.) t woo per ev seed cop win #silc, bar, fy, fo agrc, orn grandis Myrtaceae T (Aus.) t woo per ev seed cop win #silc, bar, sa, fo, gr orn, agrc Hakea sericea Proteaceae ST (Aus.) t/s woo per ev seed win orn, c/b, #bar fy Hedychium coccineum Zingiberaceae NT (Asia) h her per ev seed rhz bir, wat #orn fo coronarium Zingiberaceae NT (Asia) h her per ev seed rhz bir, wat #orn fo gardnerianum Zingiberaceae NT (Asia) h her per ev seed rhz bir, wat #orn fo Ipomoea indica Convolvulaceae T (Am.) c her per ev seed ?win, wat #orn sa, fo purpurea Convolvulaceae T (Am.) c her ann germ seed ?win, wat #orn sa, fo Jacaranda mimosifolia Bignoniaceae T (Am.) t woo per d seed cop win #orn sa, fo Lantana camara Verbenaceae T (Am.) s woo per ev/d seed cop, bir #orn, bar sa, fo, gr ?run Leptospermum laevigatum Myrtaceae ST (Aus.) t/s woo per ev seed win,wat orn, #bar, c/b fy Ligustrum japonicum Oleaceae NT (Asia) t/s woo per ev seed cop bir orn, #bar fo lucidum Oleaceae NT (Asia) t/s woo per ev seed cop bir orn, #bar fo Litsea glutinosa Lauraceae T (Asia) t/s woo per ev seed ?cop bir #orn fo Macfadyena unguis-cati Bignoniaceae T (Am.) c sem per ev/d seed cop, tub win #orn sa, fo Melia azedarach Meliaceae T(Aus.) t woo per d seed cop bir, wat #orn sa, fo, gr Morus alba Moraceae NT (Asia) t woo per d seed cop bir orn, #agrc sa, fo, gr Nephrolepis exaltata Davalliaceae T (Am.) h her per ev sp sto, tub win, hum #orn fo Nicotiana glauca Solanaceae T (Am.) t/s woo per ev seed cop win, wat, #orn sa, fy, nk, sk ?hum (soil) Opuntia fi cus-indica Cactaceae T (Am.) t/s suc per ev seed div mam, bir #agrc, bar sa, fy, fo, gr, nk, sk robusta Cactaceae T (Am.) ?t/s suc per ev seed div mam, bir #agrc, bar nk stricta Cactaceae T (Am.) s suc per ev seed div mam, bir #orn sa Paraserianthes lophantha Fabaceae ST (Aus.) t/s woo per ev seed ?cop wat #orn, agrc fy Passifl ora edulis Passifl oraceae T (Am.) c her per ev seed mam, bir orn, #agrc fo Pennisetum clandestinum Poaceae T (Afr.) g her per ev/d seed rhz, sto ?win, hum #c/b, agrc fo Pereskia aculeata Cactaceae T (Am.) s/c suc per ev seed div bir, ?mam, #bar, orn fo hum

APPENDIX 6.—Characteristics of prominent invaders in study area (cont.)

Plant name Family Origin GF W LC P SR VR Disp. agent Cult. use VC Pinus patula Pinaceae T (Am.) t woo per ev seed win #silc, bar, fo, gr orn pinaster Pinaceae NT (Med.) t woo per ev seed win #silc, bar fy, fo radiata Pinaceae NT (N Am.) t woo per ev seed win #silc, bar fy, fo Populus alba Salicaceae NT (Eur., N t woo per d suc,cop wat silc, #bar, sa, fo, gr, nk Afr. & Asia) orn deltoides Salicaceae NT (N Am.) t woo per d seed suc, cop wat, win #silc, agrc, gr orn nigra var. italica Salicaceae NT (Eur. & t woo per ev/d suc, cop wat orn, #bar, gr Asia) c/b, agrc ×canescens Salicaceae NT (Eur. & t woo per ev/d suc, cop wat silc, #c/b, sa, fy, fo, gr, Asia) bar, orn nk, sk Prosopis glandulosa var. torreyana Fabaceae NT (N Am.) t/s woo per d seed cop mam, wat #agrc, orn sa, nk, sk velutina Fabaceae NT (N Am.) t/s woo per d seed cop mam, wat #agrc, orn sa, nk, sk Prunus persica Rosaceae NT. (Asia) t woo per d seed ?cop hum #agrc, orn gr Psidium guajava Myrtaceae T (Am.) t/s woo per ev seed suc, cop mam, bir, #agrc, orn sa, fo hum Pyracantha angustifolia Rosaceae N Temp. s woo per ev seed ?cop bir orn, #bar gr (Asia) coccinea Rosaceae N Temp. s woo per ev seed ?cop bir orn, #bar gr (Eur. & Asia) crenulata Rosaceae NT (Asia) s woo per ev seed ?cop bir orn, #bar gr Quercus robur Fagaceae NT (Eur. & t woo per d seed ?cop wat, ?mam #orn, agrc fy Asia) (squirrels) Ricinus communis Euphorbiaceae T (Afr.) t/s woo var ev/d seed cop wat, hum #agrc, orn sa, fy, fo, sk Robinia pseudoacacia Fabaceae NT (N Am.) t woo per d seed suc, cop wat, hum orn, #c/b, gr bar, agrc Rosa rubiginosa Rosaceae NT (Asia) s woo per d seed ?cop mam, bir #orn, bar, gr agrc Rubus cuneifolius Rosaceae NT (N Am.) s woo per ev/d seed suc, cop bir #agrc sa, fo, gr ?pascuus Rosaceae NT (N Am.) s woo per ev/d seed suc, cop bir none sa, gr fruticosus Rosaceae NT (Eur.) s woo per ev/d seed suc, cop bir #agrc sa, fy, fo, gr ×proteus Rosaceae hybrid s woo per ev/d seed suc, cop bir none sa, gr origin (N Am. × S Afr.) Salix babylonica Salicaceae NT (Asia) t woo per d div wat, hum orn, #c/b, sa, fy, fo, agrc gr, nk fragilis Salicaceae NT (Eur. & t woo per d div wat, hum orn, #c/b, gr Asia) ?agrc Salsola kali/tragus Chenopodiaceae NT (Eur. & h her ann germ seed win none nk Asia) Schinus molle Anacardiaceae T (Am.) t woo per ev seed ?cop bir #orn, bar nk, sk terebinthifolius Anacardiaceae T (Am.) t/s woo per ev seed ?cop bir orn, #bar fo Senna didymobotrya Fabaceae T (Am.) t/s woo per ev seed ?cop wat, hum orn, #bar sa, fo Sesbania punicea Fabaceae T (Am.) t/s woo per ev/d seed wat #orn sa, fy, gr Solanum elaeagnifolium Solanaceae ST (S Am.) h/s sem per stems seed rhz ?bir none nk d mauritianum Solanaceae T (Am.) t/s woo per ev seed cop bir #orn sa, fy, fo, gr pseudocapsicum Solanaceae T (Am.) h/s sem per ev seed ?cop bir #orn fo seaforthianum Solanaceae T (Am.) c her per ev/d seed cop bir #orn sa, fo Origin: ST, southern temperate, south of or straddling Tropic of Capricorn; NT, northern temperate, north of or straddling Tropic of Cancer; T, tropical, between or straddling Tropics of Capricorn and Cancer. GF, growth form: tree; tree/shrub; shrub; herb; herb/shrub; grass; grass/reed; climber; shrub/climber. W, woodiness: woody; semi-woody; herbaceous; succulent. LC, life cycle: perennial, annual, variable, biennial. P, perennation: evergreen; deciduous; evergreen/deciduous; variable; germinative. SR, sexual reproduction: seed, spore. VR, vegetative reproduction: coppice; sucker; division; rhz, rhizome; bulbil; stolon; tuber; runner. Dispersal agent: wind; water; bird; mammal; human; ant. Cultivated use: # primary (= major) use; ornamental; barrier; cover/binder; agricultural crop; silvicultural crop; none. VC, vegetation category: savanna biome; fynbos biome; forest habitats; grassland biome; nama-karoo biome; succulent karoo biome. Afr., Africa; Am., America; Aus., Australia; Eur., Europe; Med., Mediterranean. 248 Bothalia 37,2 (2007)

APPENDIX 6.—Characteristics of prominent invaders in study area (cont.)

Plant name Family Origin GF W LC P SR VR Disp. agent Cult. use VC Tamarix chinensis Tamaricaceae NT (Asia) t/s woo per ev/d seed ?cop wat, win #?orn, c/b, nk, sk agrc ramosissima Tamaricaceae NT (Eur. & t/s woo per ev/d seed cop, suc wat, win #?orn, c/b, nk, sk Asia) agrc Tithonia diversifolia Asteraceae T (Am.) s her var var seed ?win #orn sa, fo Xanthium strumarium Asteraceae T (Am.) h her ann germ seed wat, mam none sa, fo Origin: ST, southern temperate, south of or straddling Tropic of Capricorn; NT, northern temperate, north of or straddling Tropic of Cancer; T, tropical, between or straddling Tropics of Capricorn and Cancer. GF, growth form: tree; tree/shrub; shrub; herb; herb/shrub; grass; grass/reed; climber; shrub/climber. W, woodiness: woody; semi-woody; herbaceous; succulent. LC, life cycle: perennial, annual, variable, biennial. P, perennation: evergreen; deciduous; evergreen/deciduous; variable; germinative. SR, sexual reproduction: seed, spore. VR, vegetative reproduction: coppice; sucker; division; rhz, rhizome; bulbil; stolon; tuber; runner. Dispersal agent: wind; water; bird; mammal; human; ant. Cultivated use: # primary (= major) use; ornamental; barrier; cover/binder; agricultural crop; silvicultural crop; none. VC, vegetation category: savanna biome; fynbos biome; forest habitats; grassland biome; nama-karoo biome; succulent karoo biome. Afr., Africa; Am., America; Aus., Australia; Eur., Europe; Med., Mediterranean. 2006-12-12

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